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QL 
461 
.A373X 
ENT 



MEMOIRS 
OF THE 

INTOMOLOGICAL SOCIETY 
NUMBER 24 



THE HIGHER CLASSIFICATION, 

PHYLOGENY AND ZOOGEOGRAPHY 

OF THE SATYRIDAE (LEPIDOPTERA) 



BY 



LEE D. MILLER 




BL1SHED BY THE AMEBJ INTOMOLOGICAL SOCIETY 

AT THE ACADEMY OF NATURAL SCIENCES 
PHILADELPHIA 

1968 



MEMOIRS 

OF THE 

AMERICAN ENTOMOLOGICAL SOCIETY 

NUMBER 24 



THE HIGHER CLASSIFICATION, 
PHYLOGENY AND ZOOGEOGRAPHY 
OF THE SATYRIDAE (LEPIDOPTERA) 



BY 



LEE D. MILLER 




PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY 

AT THE ACADEMY OF NATURAL SCIENCES 

PHILADELPHIA 

1968 




Selwyn S. Roback 
Editor 



(Issued October 22, 1968) 



Printed in the United States of America 



To the memory of Dr. Richard M. Fox 

— Teacher, inspiration, colleague, friend. 

He is and will be missed. 



Preface 

No work is ever truly done alone: this paper is ample proof of 
the statement. I wish to take this opportunity to thank the many 
people and organizations who have helped to make it a reality. 

The research was supported in part by National Science Founda- 
tion grant number GB-2928 (R. M. Fox, Principal Investigator). 
The National Science Foundation, through Office of Science Infor- 
mation Services, made possible the publication of this work by grant 
number GN-652. 

Thanks are due the following individuals and institutions for their 
aid, loan of materials, use of facilities and general encouragement: 
Carnegie Museum, particularly Dr. M. Graham Netting, Director, 
Drs. Richard M. Fox and George E. Wallace and Mr. Harry K. 
Clench of the Section of Insects and Spiders; the Department of En- 
tomology, British Museum (Natural History), especially Messrs. J. P. 
Doncaster, Keeper of Entomology, and N. D. Riley, P. E. S. Whal- 
ley, T. G. Howarth and M. Clifton of the Lepidoptera section; and 
the Departments of Biology of the University of Pittsburgh and The 
Catholic University of America. Without their cooperation this pro- 
ject could not have been completed. 

The following individuals read the original manuscript which was 
submitted in quite different form as a dissertation at the University of 
Pittsburgh: Drs. Fox, Peter Gray, Eliot B. Spiess, Malcolm T. Jollie 
and Kenneth W. Cummins, Department of Biology, University of 
Pittsburgh, C. C. Li, Department of Biostatistics, Graduate School of 
Public Health. University of Pittsburgh, Craig C. Black and Mary 
Dawson, Section of Vertebrate Paleontology, Carnegie Museum. All 
were available for consultation and deserve great thanks. 

Mr. Clifton read the part of the manuscript dealing with the 
Haeterinae and made the necessary revisions on it based upon his 
forthcoming revision. Dr. Norman Tindale of the South Australian 
Museum read and commented upon the zoogeographic analyses, espe- 
cially those dealing with Australia. Dr. F. Martin Brown and Mr. 
Harry K. Clench read all or part of the manuscript and commented 
upon it. Particular thanks, however, are due Dr. Fox, who read and 
carefully critiqued the entire manuscript. He offered countless valu- 
able suggestions, most of which have been incorporated into the manu- 
script. 



Preparation of the manuscript and the figures has been aided 
greatly through the activities of two of my graduate students, Misses 
Jacqueline Y. Sessi and Dorothy A. Shea. 

Finally, thanks are due the many unnamed friends and colleagues 
who, knowingly or not, have contributed ideas and moral support for 
this undertaking. 



Table of Contents 

Page 

INTRODUCTION 1 

The Problem 1 

Historical Sketch 1 

Material Studied and Methods Employed 4 

COMPARATIVE MORPHOLOGY 7 

Antennae 7 

Head 7 

Labial Palpi 7 

Thorax 8 

Forelegs 8 

Male forelegs 8 

Female forelegs 9 

Walking Legs 10 

Wings 10 

Forewings 1 1 

Hindwings 12 

Androconial patches 13 

Male Genitalia 14 

SYSTEMATIC REVISION OF THE GENERA OF THE SATYRIDAE 14 

Family Satyridae Boisduval 15 

Key to the Subfamilies of the Satyridae 18 

Subfamily Haeterinae Herrich-Schaffer 19 

Tribe Haeterini Herrich-Schaffer 20 

Genera Included in the Haeterini 22 

Subfamily Brassolinae Boisduval 23 

Tribe Brassolini Boisduval 23 

Genera Included in the Brassolini 27 

Subfamily Biinae Herrich-Schaffer 28 

Key to the Tribes of the Biinae 29 

Tribe Antirrhini, new tribe 29 

Genera Included in the Antirrhini 31 

Tribe Biini Herrich-Schaffer 32 

Genus Included in the Biini 34 

Tribe Melanitini, new tribe 34 

Genera Included in the Melanitini 36 

Genus Manataria Kirby 36 

Subfamily Elymniinae Herrich-Schaffer 38 

Key to the Tribes of the Elymniinae 40 

Tribe Lethini Clark 40 

Genera Included in the Lethini 47 



MEM. AMER. ENT. SOC, 24 



Tribe Zetherini, new tribe 52 

Genera Included in the Zetherini 53 

Tribe Elymniini Herrich-Schaffer 54 

Genera Included in the Elymniini 56 

Tribe Mycalesini, new tribe 57 

Genera Included in the Mycalesini 60 

Subfamily Eritinae, new subfamily 64 

Tribe Eritini, new tribe 64 

Genera Included in the Eritini 66 

Subfamily Ragadiinae Herrich-Schaffer 67 

Tribe Ragadiini Herrich-Schaffer 67 

Genera Included in the Ragadiini 69 

Subfamily Satyrinae Boisduval 69 

Key to the Tribes of the Satyrinae 73 

Tribe Hypocystini, new tribe 74 

Genera Included in the Hypocystini 80 

Tribe Ypthimini, new tribe 81 

Genera Included in the Ypthimini 85 

Genus Palaeonympha Butler 87 

Tribe Euptychiini, new tribe 89 

Genera Included in the Euptychiini 92 

Tribe Coenonymphini, new tribe 95 

Genera Included in the Coenonymphini 98 

Tribe Maniolini Hampson 98 

Genera Included in the Maniolini 101 

Tribe Erebiini Tutt 101 

Genera Included in the Erebiini 104 

Tribe Dirini, new tribe 105 

Genera Included in the Dirini 108 

Tribe Pronophilini Clark 108 

Genera Included in the Pronophilini 114 

Tribe Satyrini Boisduval 119 

Genera Included in the Satyrini 123 

Tribe Melanargiini Verity 124 

Genera Included in the Melanargiini 126 

Genera of Uncertain Position 127 

Genus Pamperis Heimlich 127 

Genus Setodocis Billberg 128 



u 



THE EVOLUTION AND ZOOGEOGRAPHY OF THE SATYRIDAE 128 

Origin of the Satyridae 131 

The Later History of the Satyridae 134 

Derivation of the Satyrid Faunae of the World 142 

The Neotropics 145 

The Nearctic 145 

The Palearctic 147 

The Indo-Malayan Region 148 

The Australian Region, including the Pacific Islands 149 

The Ethiopian Region 150 

Madagascar 152 

BIBLIOGRAPHY 153 

INDEX 164 



MEM. AMER. ENT. SOC, 24 

iii 



Memoirs 

of THE 

American Entomological Society 

Number 24 



THE HIGHER CLASSIFICATION, 

PHYLOGENY AND zoogeography OF 

THE SATYRIDAE (LEPIDOPTERA) 

By 
Lee D. Miller 

Introduction 

The Problem. — The objective of this study, briefly, is the anal- 
ysis, both in space and, where possible, in time, of the evolution, phy- 
logeny and zoogeography of the higher taxa of the Satyridae. The 
existing studies on the family have been generally regional, and little 
progress has been made on interrelating the faunae of the various re- 
gions of the world. Furthermore, these studies were not based on 
modern, multi-variant taxonomic principles; hence, as will be shown, 
some fundamental errors in relationships have been perpetrated. Since 
no adequate taxonomic revision of the higher categories of the Saty- 
ridae is available, it has been necessary to complete one as a basis for 
the evolutionary and zoogeographic discussions. Using morphologi- 
cal and distributional data as bases, a scheme for the evolution and 
zoogeography of the Satyridae is proposed, and finally a phylogeny of 
the family is projected. 

Historical Sketch. — Linne considered all butterflies congen- 
eric. In the Tenth Edition of his Sy sterna Naturae (1758), the start- 
ing point of zoological nomenclature, he published descriptions of a 

MEM. AMER. ENT. SOC, 24 



THE SATYRIDAE 



number of satyrids, placing them with all other butterflies in the genus 
Papilio. The first nomenclatorially valid satyrid genus, Maniola, was 
erected by Schrank (1801) in his Fauna Boica. Fabricius (1807) 
described several more genera in Illiger's rare Magazin fur Inseckt- 
enkunde. Latreille (1810) proposed Satyrus in his Considerations 
generates sur Yordre natural des animaux . . . crustaces . . . arach- 
nides . . . et insectes . . . Hiibner, between the years 1806 and 
1819 (the latter date is bibliographically indeterminate and is brack- 
eted in all succeeding references), published many satyrid generic 
names in the Sammlung Exotischer Schmetterlinge and the Vcrzeich- 
niss bekannter Schmettlinge [sic]. Several other authors, notably 
Billberg (1820) and Meigen (1829), proposed early generic names 
for satyrids, but it was not until 1836 that the family was described, 
as "Satyrides", by Boisduval. That author also first proposed the 
Brassolinae, as "Brassolides". 

The first comprehensive work devoted to the butterflies of the 
world in which the treatment was anything like a modern one was 
The genera of diurnal Lepidoptera by E. Doubleday, J. O. Westwood 
and W. C. Hewitson which appeared in two volumes between 1846 
and 1852. In this work the Satyridae were defined, the brassolines 
delimited (as a subfamily of the Morphidae) and many generic names 
were added to the Satyridae. The practice of separating the brasso- 
lines from the rest of the satyrids became firmly established with this 
publication. The majority of the satyrid generic names were de- 
scribed in this work between 1849 and 1851, after Doubleday's death, 
and are usually considered to have been authored by Westwood. 
The authors always referred to this work as "Doubleday, Westwood 
and Hewitson", even though Hewitsons chief role was as illustrator; 
but Brown (1941) holds that text descriptions must be credited to 
either Doubleday or Westwood alone, while names first proposed in 
plates (the text and the plates were often separately published) should 
be credited to Doubleday and Hewitson or to Westwood and Hewit- 
son. Accordingly, in those instances where the text appeared simul- 
taneously with, or prior to, the plates, I am crediting authorship to 
either Doubleday or Westwood, but if the plates appeared before the 
text descriptions, credit must be given to Doubleday and Hewitson or 
to Westwood and Hewitson. 

The first attempt at a higher classification of the satyrids ( actually, 



LEE D. MILLER 3 

of all the butterflies) was that of Herrich-Schaffer (1864). For the 
first time the satyrids were split beyond the Satyrinae and Brassolinae, 
as "Satyrina" and "Brassolina". The Biinae ("Biina"), Haeterinae 
("Hetaerina" [sic]), Ragadiinae ("Ragadiina") and Elymniinae 
("Elymniina") were established. This work has been too long over- 
looked by systematists — all but one of the presently recognized saty- 
rid subfamilies were defined in it. Rober (1892), in the second vol- 
ume of Staudinger and Schatz' Exotische Schmetterlinge, divided the 
satyrid genera into six groups with a residue of indeterminate genera. 
Rober's groups generally correspond to major divisions of the Saty- 
ridae, but they were not nomenclatorially designated. Moore in the 
first and second volumes of his Lepidoptera Indica, published between 
1890 and 1893, proposed a great many generic names but considered 
these genera to be apportioned into only the Satyrinae and Elym- 
niinae. These same divisions are to be found in Gaede ( 193 1 ) . The 
Rober scheme was followed with little modification by Fruhstorfer, 
Weymer and Aurivillius in their respective sections of Seitz' monu- 
mental Die Grossschmetterlinge der Erde. 

More recently Clark (1947) subdivided the satyrids, exclusive of 
the brassolines, into the Satyrinae, Enodiinae, Pronophilinae, Elym- 
niinae and Pierellinae (a synonym of Haeterinae). In 1948 he 
emended Enodiinae to Lethinae and divided the brassolines, as a sep- 
arate family, into the Brassolinae, Caliginae and Biinae (this last not 
properly a brassoline). In neither paper did Clark give definitive 
reasons for his classification, hence he has been criticized by such 
authors as Ehrlich (1958) who proposed a higher classification of 
the "true butterflies" (Papilionoidea), retaining the separation of the 
brassolines and the satyrids and placing the former with the Mor- 
phidae. 

Table 1 attempts to relate the classifications of Herrich-Schaffer 
(1864) and Clark (1947, 1948) with that proposed in the present 
paper. In many instances particular genera were not apportioned by 
either Herrich-Schaffer or Clark, and those authors' intentions can 
only be guessed. 

With the exception of Clark (1947, 1948) and Ehrlich (1958), 
no modern author has attempted to place the satyrids in their proper 
evolutionary positions, and there has been no zoogeographic analysis 
of this family. Schwanwitsch (1924), however, relied heavily on 

MEM. AMER. ENT. SOC, 24 



THE SATYRIDAE 



Table 1 



A comparison of the present subfamilial classification of the Satyridae with 
those proposed by Herrich-Schaffer (1864) and Clark (1947, 1948). 

Present Herrich-Schaffer's Clark's 

classification classification classification 

Haeterinae Hetaerina (sic) Pierellinae 

Brassolinae Brassolina Brassolinae 

Caliginae 

Biinae Biina Biinae (of Brassolidae) 

Satyrina (part) Satyrinae (part) 

Elymniinae Elymniina Elymniinae 

Satyrina (part) Enodiinae 

(=Lethinae) 

Eritinae Satyrina Satyrinae 

Ragadiinae Ragadiina Satyrinae 

Satyrinae Satyrina Satyrinae (part) 

Pronophilinae 

satyrids in his study of the evolutionary trends toward the develop- 
ment of the "basic nymphalid pattern". This work was followed by 
a number of other papers concerning pattern modifications, two of 
which are of interest here: in 1925 he published an analysis of the 
Pierella-type pattern, a paper which has since become a classic in the 
explanation of the morphological migration of pattern elements; and 
in 1931 he analyzed the highly modified Melanargia-type pattern. 
Both papers were significant, but the one explaining "pierellization" 
has had the most far-reaching implications. 

Material Studied and Methods Employed. — Since there are 
between 2500 and 3000 species in the Satyridae, it would be im- 
practical to attempt a complete revision of all the species in a single 
work. The number of described genera, on the other hand, is man- 
ageable — fewer than 400. 

A fair axiom in systematics may be stated more or less as fol- 
lows: systematic decisions at one taxonomic level should be reached 
by examination of most (preferably all) of the entities at the next 
lower major taxonomic level. In other words, if one is revising a 
genus, one should see representatives of the species and subspecies 
included in it to make meaningful taxonomic judgments at the spe- 
cific and generic levels. In the present study examples of the already- 
named genera were examined, hence, taxonomic judgments are made 
at the tribal level, and no synonymization of nomenclatorially valid 



LEE D. MILLER 5 

genera (only pure objective synonyms) has been attempted, nor have 
the several new genera which certainly exist been named. This is 
work for careful species-by-species revisions. It is certain that many 
of the nomenclatorially valid genera are not biological entities; for 
example, I am unable to ascertain any significant differences at the 
generic level between the seven nomenclatorially valid genera included 
in the satyrine tribe Melanargiini. Nevertheless, all of the nomencla- 
torially valid names are carried in this revision. 

For the initial work on this project the collection of Carnegie 
Museum was examined for representatives of the known genera. It 
soon was apparent that the material at hand was not sufficient, so 
two months were spent at that Mecca of butterfly systematists, the 
British Museum (Natural History), examining those genera which 
were either unrepresented at Carnegie Museum or only poorly rep- 
resented. Material was found and examined in these collections rep- 
resenting all but four of the genera that had come to my attention by 
August, 1964. Those genera that have been described since that 
time are listed in what I believe to be their proper systematic posi- 
tions, but none of them has been examined as critically as have the 
others. 

Sokal and Sneath (1963: 161-162) explain the "exemplar meth- 
od" of taxonomy as follows: a taxon may be characterized by a small 
number of entities within it (even a single specimen) because the 
variation within taxa is considered to be less than that between taxa. 
This is somewhat the method that has been employed in the present 
study. The type-species of a genus is considered to be typical of it, 
and the various genera are defined chiefly by their type-species in the 
following pages. Occasionally the type-species is aberrant, but it is 
the type-species which bears nomenclatorial "responsibility" for the 
generic name, in any event. Where possible, other species of the 
genera were examined, though not in as great detail as the type- 
species. 

The systematic revision is based entirely upon external morpho- 
logical characteristics with many attributes being considered. Wing 
venation drawings were made from specimens either with an ocular 
grid under a dissecting microscope or with a projection device for 
larger specimens. Antennae, palpi, fore-, mid- and hindlegs were re- 
moved from the specimen, bleached in clorox and mounted on perma- 

MEM. AMER. ENT. SOC, 24 



b THE SATYRIDAE 

nent slides. Drawings of these structures were made with a camera 
lucida. 

The taxonomic and phylogenetic analysis of the family has been 
generally based on the premises that ( 1 ) entities which more nearly 
resemble one another in many characteristics are most closely re- 
lated, ( 2 ) structures which are reduced and fused represent advanced 
conditions (Fox, 1956: 24; Fox and Fox, 1964: 71-72; Brown, 
1965) and (3) those organisms which bear advanced characteristics 
(not just a single advanced character, but several of them) are prob- 
ably themselves advanced (for example, see Mayr, 1965). This is 
in accord with the current paleontological evidence drawn from ex- 
isting fossil records. 

The only taxonomically valuable structures not utilized in this 
study were the male and female genitalia. Examination of several 
regional works (for example, Hayward, 1953; Forster, 1964) shows 
little reliance can be placed on genitalic structures for classification 
above the generic level, although these structures are certainly of great 
importance in specific, and sometimes generic, determination. 

Authorship of higher taxa has caused some problems. According 
to Art. 36 of the International Code of Zoological Nomenclature 
(1961), the proposition of the family Satyridae by Boisduval (1836, 
as "Satyrides") implicitly creates also the subfamily Satyrinae and 
the tribe Satyrini. It is this rule which has been followed through- 
out the present paper for the assignment of authorship to the higher 
taxa. 

The zoogeographic analysis has been carried on in light of the 
principles of vertebrate zoogeography, which is firmly based on and 
supported by fossil evidence. For a summary of such evidence see 
Darlington (1957). There are too few butterfly fossils to support or 
refute any zoogeographic statement. Heavy reliance has been placed 
on the presence in one area or another of relict, annectant forms: 
such species may occur nearer the place of origin of the taxon than 
do others. Although fossil evidence shows that such groups as horses 
and rhinoceri evolved more extensively in the Nearctic than in the 
Palearctic, yet are now absent from the former, many other groups 
have evolved more or less in the places of their greatest present-day 
density and diversity. In the absence of fossil evidence to the con- 
trary, the latter situation is considered to be likely in the satyrid but- 
terflies. 



LEE D. MILLER 



Comparative Morphology 



The characters utilized in this study are all external and mor- 
phological. Their distribution and evolutionary patterns are given 
in the following pages. 

Antennae. — Four measurements were made on the antennae: 
the total length; the length of the club, where the club is a discreet 
entity; the width of the shaft and the width of the club at its thickest 
point. The antennae of Satyridae demonstrate the tricarinate con- 
dition typical of the Nymphaloidea (Fig. 249), although the ridges 
may not be apparent throughout the entire length of the antennae. 
The antennae are scaled, but only on the basal few segments in the 
Haeterinae. In general, the antennal club is not greatly thickened 
nor flattened at the tip in the primitive satyrids (Figs. 114, 122, 128 
which are some of the best developed ones), thereby following the 
general primitive condition in butterflies. Many of the more advanced 
Satyrinae. conversely, have the antennal club strongly developed, flat- 
tened at the tips and occasionally exhibiting bizarre configurations in 
some Hypocystini and Pronophilini (Figs. 163, 166, 169, 291, for 
example ) . The antennal clubs of other Satyrinae are shown in Figs. 
142, 151, 191, 213, 220, 235, 249, 262, 297. 

Head. — Three measurements were made on the head region: 
least intraocular distance; height of the eye and width of the eye. 
The eyes of all satyrids are entire. Little significance could be gleaned 
from these measurements, but the eyes of the Satyrinae appear to be 
more ovoid than those of other subfamilies. 

Labial Palpi. — In common with other butterflies the labial palpi 
of the Satyridae are three-segmented. The basal segment is so firmly 
attached to the head that it was often broken in dissection, hence is 
little used in the present study. Five measurements were made on 
the palpi: the length of the first segment, when it was not broken; the 
length of the second segment; the length of the third segment; the 
maximum width of the second segment and the maximum length of 
the hairs of the second segment. The relationship between the lengths 
of the second and third segments of the palpi varies, often without 
regard to phyletic groups. For example, most of the Satyrini display 
an extremely short third palpal segment, about one-fifth as long as 
the second, whereas the Pronophilini. which are closely related to the 
Satyrini and may have been derived from a fairly recent common an- 

MEM. AMER. ENT. SOC, 24 



8 THE SATYRIDAE 

cestor, show the longest third palpal segment of any satyrid tribe — 
in some instances over half as long as the second segment. The sec- 
ond segment is generally between five and eight times as long as it is 
wide, again with no apparent phylogenetic correlation. The relative 
lengths of the hairs of the second segment of the palpi show some in- 
teresting trends. In general the hairs are longer in the Satyrinae than 
in the other subfamilies, but this tendency is modified in those groups 
which have both tropical and temperate or alpine members. In such 
instances the relative length of the hairs increases as the altitude or 
latitude increases. Examples of the labial palpi of the Satyridae are 
shown in many figures in the text, such as Figs. 3, 10, 22, 30, 35, 50, 
82, 93, 100, 129, 135, 143, 172, 198, 214, 225, 236, 250, 263, 
298, 311. 

Thorax. — Only one thoracic measurement was made: the length 
of the dorsal thoracic elements. This measurement, unfortunately, is 
generally dependent only upon the absolute size of the insect. 

Forelegs. — The forelegs of both sexes are more or less reduced 
in all satyrids, as in other members of the Nymphaloidea. The fore- 
coxae are moveable, as determined by examination of many (though 
not all ) genera. The forecoxae are not reduced to the extent that the 
more distal elements of the forelegs are. The reduction of the pro- 
thoracic femur, tibia and tarsus has long been the criterion for re- 
garding the Nymphaloidea as the most advanced of the lepidopterous 
insects, a concept recently challenged by Ehrlich (1958). 

Male forelegs. — Four measurements were made on the male 
forelegs: the length of the tarsus; the length of the first tarsal sub- 
segment; the length of the tibia and the length of the femur. The 
miniaturization of the male forelegs generally follows phylogenetic 
lines, being least miniaturized (and reduced; for a discussion of the 
terms "miniaturized" and "reduced" see Fox, 1967) in the Haeter- 
inae, Brassolinae, Biinae, Elymniinae and Eritinae and most in the 
Ragadiinae and Satyrinae (compare the graphs in Figs. 1, 20, 170, 
309, for example). In the first five subfamilies the femur, tibia and 
tarsus usually are all relatively well developed (an exception is Cae- 
rois, Fig. 27, where the tibia and the tarsus are much miniaturized). 
The foretarsus in these primitive subfamilies is typically monomerous, 
but some genera may have up to four freely articulating subsegments. 
Examples of the forelegs of typical primitive satyrids are Figs. 4, 11, 



LEE D. MILLER V 

23, 51, etc. In the Ragadiinae and Satyrinae the forelegs are much 
more miniaturized and reduced than in the other five subfamilies, and 
occasionally there is an apparent total loss of the tarsus (Ypthimini, 
Fig. 177 ). The tibia and femur are also greatly miniaturized, but the 
coxa does not seem affected by this general trend. An anomalous 
situation exists in the Satyrini and Erebiini, however, where a few 
genera display great miniaturization of the total foreleg but still re- 
tain up to four distinct, though tiny, tarsal subsegments. The male 
foreleg of Ragadia is shown in Fig. 136, and those of representative 
Satyrinae are shown in Figs. 144, 173, 193, 199,215,226,237,251, 
266, 299, 312, etc. 

Female forelegs. — Five measurements were made on the female 
forelegs: the length of the tarsus; the length of the first tarsal sub- 
segment; the length of the fifth tarsal subsegment, if any was present; 
the length of the tibia and the length of the femur. The female fore- 
legs potentially show more precise inter- and intrasubfamilial rela- 
tionships than do those of the males. The forelegs of the Haeterinae, 
Brassolinae, Biinae and Elymniinae are quite well developed (Figs. 1, 
20, 41, 48, 88, 89, 98), always have pentamerous tarsi and usually 
have posttarsal pulvilli present, except a few of the more advanced 
Elymniinae. In these families the tarsus is not, or only weakly, 
clubbed (for example, Figs. 5, 15, 24, 52, 112). In only two genera, 
Paralethe and Antirrhea (Fig. 24) are there posttarsal claws, and 
always there is but a single claw. This represents a more advanced 
condition generally than that shown in the Ithomiidae (Fox, 1956: 
13). The female forelegs of the Eritinae (Figs. 125, 131) and the 
Ragadiinae (Fig. 137) are miniaturized beyond the point shown in 
the preceding subfamilies, but they retain the full complement of 
tarsal subsegments; the tarsus itself is clubbed and not so freely ar- 
ticulated as in the primitive subfamilies. In the Satyrinae the fore- 
legs range from miniaturized, but with five tarsal subsegments, to 
highly reduced, as much so or more than in the males. Hypocystini, 
Ypthimini (interestingly where the greatest reduction takes place in 
the male foreleg), Euptychiini, most Pronophilini and some members 
of several other tribes have pentamerous tarsi (Figs. 145, 174, 194, 
200, 267, for example), but the tarsus is always clubbed. The fore- 
legs of female Satyrini, Melanargiini, Dirini, Coenonymphini and 
Maniolini are always reduced and apparently lack one or more tarsal 

MEM. AMER. ENT. SOC, 24 



10 THE SATYRIDAE 

subsegments (Figs. 216, 227, 238, 252, 300, 313, for example). 
The miniaturization of the forelegs of Satyrinae through the phyletic 
lines may be seen by examination of Figs. 140, 170, 211, 295, 309, 
etc. 

Walking Legs. — Six measurements were made on both the 
mesothoracic and metathoracic legs: the length of the tarsal claws; 
the length of the tarsus, exclusive of the claws; the length of the first 
tarsal subsegment; the length of the tibia; the length of the tibial spurs, 
if any, and the length of the femur. The more important taxonomic 
characters are on the mesothoracic legs. The lack of tibial spurs 
characterizes the Haeterinae, Biini and Elymniini (Figs. 6, 33, 96), 
and these spurs have been lost sporadically in such other groups as 
the Ypthimini (Fig. 179) and Coenonymphini (Fig. 221). The 
posttarsal claws are bifid in the Melanitini (Fig. 39) and in a few 
other genera (Samanta and Manataria, Fig. 47). In all other Saty- 
ridae the claws are simple. The relative lengths of the midtibia and 
the first midtarsal subsegment are important taxonomically, particu- 
larly within the Satyrinae. Usually the midtibia is between one and 
a half and two times as long as the first midtarsal subsegment, but 
the midtibia is barely longer than the first tarsal subsegment in the 
Satyrini (Fig. 301) and in some Dirini (Fig. 259), whereas the mid- 
tibia is always more than twice as long as the basal midtarsal sub- 
segment in the Coenonymphini (Fig. 217). All members of the 
Satyrus series of the Satyrini display a heavy spine at the dorsal distal 
end of the midtibia (Fig. 301), a characteristic weakly shown by 
some Dirini (as Torynesis, Fig. 259), thereby indicating a close re- 
lationship between these two tribes. Dorsal spinosity of the midtibia 
is shown by some members of most groups and is characteristic of 
others: Haeterinae, Zetherini, Pronophilini, Dirini, Satyrini and Me- 
lanargiini. 

Examination of the charts of relative lengths of legs shows that 
the hindlegs are occasionally significantly longer (Satyrini, Fig. 295, 
for example) or shorter (the Pronophila series of the Pronophilini, 
Fig. 260, for example) than the midlegs, but generally these legs are 
subequal. 

Wings. — The wings have long been the most-used basis for the 
classification of butterflies, both from the aspect of venation and the 
distribution of androconial patches. Undoubtedly too much empha- 



LEE D. MILLER 1 I 

sis has been placed in the past on wing venation and androconial dis- 
tribution alone, but it is illogical to ignore the structure of the wings 
because of previous preoccupation. 

Forewings: The following four measurements were made to de- 
limit the shape of the forewing: the length from the base to the end 
of vein Rr,; the length from the base to the end of M2; the length from 
the base to the end of 2A and the distance between the end of R-, and 
the end of 2 A. The cell was delimited by two measurements: from 
the base to the origins of Mi and M3. Five measurements were made 
to determine the relative positions of the origins of the veins: the dis- 
tance from the origin of Rs to the origin of Mi; from the origin of 
Mi to the origin of M2; from the origin of M-> to the origin of M3; 
from the origin of Ms to the origin of Cui and from the origin of Cui 
to the origin of Cul>. Inasmuch as the inflation of the forewing veins 
has long been considered a key characteristic of the Satyridae, the 
following three measurements were made: the thickest part of Sc; the 
thickest part of the cubital stem and the thickest part of 2A. In gen- 
eral the longest forewing cell is displayed by the most primitive sub- 
families, the Haeterinae (Figs. 2, 7, 8), Brassolinae (Figs. 9, 13. 14, 
17, 19) and Biinae (Figs. 21, 26, 28, 29, 34, 40), where the cell is 
frequently more than three-fifths the length of the wing. In most 
other groups the cell is about half as long as the wing, but the Elym- 
niini are characterized by an extremely short forewing cell (Figs. 90- 
92), often less than a third the length of the wing. There appear to 
be no broad phylogenetic implications shown by the relative place- 
ment of the veins along the cell, though several relationships are shown 
within subfamilies, particularly the proximity of the origins of M3 
and Cui in the Elymniini (Figs. 90-92). The classic definition of 
the Satyridae includes a statement about the inflation of the stems of 
the three basic forewing veins. This is simply not diagnostic: the 
inflation of the veins of Melanargia (Fig. 310), for example, is con- 
siderably less than that of the true nymphalid genus Mestra. Within 
the Satyridae particular inflation patterns are characteristic: the My- 
calesini are characterized by the inflation of all three stems, that of 
2A being more or less quadrate (Figs. 99, 104-106, 109, 113, 115, 
119); all stems are inflated in the Hypocystini (Figs. 141, 147-150, 
156, 161, 162, 165, 167, 168) and Coenonymphini (Figs. 212, 219), 
but the inflation of 2A is rounded, not quadrate. In contrast, most 

MEM. AMER. ENT. SOC, 24 



12 THE SATYRIDAE 

Biinae, Ragadiinae, Eritinae and Elymniinae (except the before- 
mentioned Mycalesini ) have relatively uninflated cubital stems and 
2A (Figs. 21, 29, 34, 49, 81, 90, 127, 134, for example). The pres- 
ence of a vestige of forewing vein 3A is characteristic of the Haeteri- 
nae (Figs. 2, 7, 8) and seems to connect them with the Ithomiidae. 

Hindwings. — Five measurements were made to delineate the 
shape of the hindwing and account for the majority of "tails": the 
length from the base to the end of vein Rs; the length from the base 
to the end of M :j ; the length from the base to the end of Cu 2 ; the 
length from the base to the end of 2A and the distance from the end 
of Rs to the end of 2 A. The cell was delimited by two measure- 
ments: from the base to the origins of Mi and Ms. Five measure- 
ments showed the relative placement of the veins along the cell: from 
the origin of Rs to the origin of Mi; from the origin of Mi to the 
origin of M 2 ; from the origin of Mi> to the origin of Ms; from the 
origin of Ma to the origin of Cui and from the origin of Cui to 
the origin of C\x-i. In all Satyrinae except Dirini (Figs. 248, 255, 
256) the hindwing cell averaged greater than half the length of the 
wing measured to the end of M :i (for example, Figs. 141. 171, 197, 
212, 224, 234, 261, 296, 310), whereas in the other subfamilies the 
hindwing cell is half or less the length of the wing, again particularly 
short in the Elymniini (Figs. 90-92). The actual shape of the cell, 
which is not reflected in the raw measurements, has some phyletic 
significance. The cell is distally rounded in the Haeterinae (Figs. 2, 
7, 8), perhaps representing a more generalized condition. In the 
Biinae a gradation may be seen between a rounded cell and one pro- 
duced at the origins of M 3 and Cui (Figs. 21, 26, 28, 29, 34, 40). 
The Elymniinae and the "Ypthimini complex" of the Satyrinae, as well 
as the Eritinae, have the cell produced at the origin of Ms (Figs. 49, 
90, 99, 127, 171, 197, for example), whereas the Hypocysta series 
of the Hypocystini and the "Satyrini complex" of the Satyrinae, in 
general, display a migration of crossvein mrm,! outward at M2 and a 
corresponding blunting of the cell (Figs. 141, 261, 296, 304, for ex- 
ample). 

The Ragadiinae (Figs. 134, 139) do not fit another part of the 
classic definition of the Satyridae — the closure of the cells by tubu- 
lar veins. In males of this subfamily the hindwing cell is closed by 
vestigial, non-tubular veins, apparently only for the support of an 



LEE D. MILLER 13 

androconial patch; some females have the cell completely open. 

Because of the migration of crossvein rao-m3, M2 arises nearer to 
the origin of Ms than to the origin of Mi in some satyrine tribes; in 
other groups M2 arises somewhat to much nearer Mi. Connate veins 
are generally considered an advanced characteristic of the Lepidop- 
tera, but in the Satyridae the opposite may be true. Hindwing veins 
Ms and Cui are frequently connate in the more primitive subfamilies, 
particularly the Haeterinae and the Elymniinae (for example, Figs. 
2, 60, 90), but these veins are well separated in the Satyrinae (for 
example, Figs. 261, 296), which are more advanced from a number 
of other viewpoints, as detailed in the preceding pages. This situa- 
tion is emphasized chiefly as an illustration of the independence of 
characters and the danger of reliance on too few attributes, as has 
been suggested by some authors (for example, Warren, 1947). Hind- 
wing vein Ri of the hindwing characteristically is basally free in the 
Brassolinae (Figs. 9, 13, 14, 17, 19) and Elymniini (Figs. 90-92). 

Androconial Patches. — As noted previously the distribution of 
androconial patches has been used extensively in the establishment of 
genera (particularly by Moore, 1890-1893) — probably to too great 
an extent. A few general statements may be made concerning the 
distribution of these patches. There are several sites of the produc- 
tion of androconia on the forewing, the most common of which is the 
discal area just outside the cell from the inner margin to the origin 
of Ms. This patch may be highly modified to include all or any part 
of the area and generally consists of a patch of mealy scales. Hair 
patches are less frequent on the forewing and are usually restricted 
to the area below the cubital stem on the upper surface or along 2A 
on the under surface, particularly in the Biinae and Mycalesini; re- 
sultant distortion of the venation may occur in these groups. Several 
areas generate androconial patches on the hindwing. Mealy patches 
are generally found near the end of the cell in the area of M3 (for 
example, Fig. 54). Hair patches are found in three general areas: 
above the cell near the origins of Rs and Mi (for example, Fig. 73), 
within the cell proper (particularly characteristic of the Elymniini, 
Figs. 90-92, and Ragadiinae, Figs. 134, 139) and along the anal 
veins (for example, Fig. 121). Since androconia appear sporadi- 
cally within most groups they seem to have little phylogenetic signifi- 

MEM. AMER. ENT. SOC, 24 



14 THE SATYRIDAE 

cance, though particular configurations may be characteristic of a few 
tribes. 

Male Genitalia. — Although these structures have been used ex- 
tensively by some authors for the establishment of genera (particu- 
larly by Forster. 1964), the genitalia are not considered in the pres- 
ent study because of the great variability shown in such genera as 
have not been split on genitalic grounds. While the genitalia are 
undoubtedly indispensable for the determination of intrageneric affin- 
ities, their unrestricted use as the basis for establishing genera is un- 
tenable, partly because doing so creates hierarchical difficulties in 
that genitalia often do not correlate with other structures, resulting 
in numerous monotypic genera. Too many monobasic taxa tend to 
obscure relationships within a larger group. 

Systematic Revision of the Genera of the Satyridae 

Before meaningful statements may be made concerning evolution, 
phylogeny and zoogeography, it is necessary to analyze relationships. 
No usable taxonomy presently exists for the Satyridae; the attempt in 
the following pages to apportion genera to subfamilies and tribes is 
designed to fill this need. This classification is based on many char- 
acters in the hope that it may more nearly approximate a "natural" 
classification than have some schemes in the past which were based 
on one or very few characteristics. 

It is the purpose of this revision to define, as closely as possible, 
the subfamilies and tribes which comprise the Satyridae, as well as to 
define the family itself. No single character can or should be ex- 
pected to be diagnostic of an entire group: evolution modifies char- 
acters indiscriminately, without regard to "sacred" taxonomic ones. 
That a single character — or even several — is atypical of the group 
in some genera is no reason to assign such genera to other tribes 
(Simpson, 1961a: 54-63). The practice of "splitting' across natural 
boundaries comes from the indiscriminate analysis of too few char- 
acters. 

Two other nymphaloid families are considered to be systemati- 
cally and phylogenetically close to the Satyridae: the Morphidae and 
the Ithomiidae. 

The Morphidae, which are represented in both the Old and New 
World tropics, have been regarded by many authors (e.g., Ehrlich, 
1958) to be very near the Satyridae. With few exceptions the hind- 



LEE D. MILLER 15 

wing cell of the morphids is open between veins M- and M :i , whereas 
only satyrids of the subfamily Ragadiinae have the cell closed by 
other than fully functional tubular veins. The structure of the mor- 
phid legs is substantially that shown by the primitive satyrids, but 
larval foodplant preference will separate some morphids. All the 
Indo-Australian morphids feed as larvae on monocotyledons, as do 
the satyrids, but the Neotropical members feed predominantly on di- 
cotyledons. There are a few nymphalid genera that utilize mono- 
cotyledons, but Morpho is unique among nymphaloids in commonly 
using both monocots and dicots (Ehrlich and Raven, 1965). On the 
larval foodplant preference and on pattern morphology (see Schwan- 
witsch, 1924) the New World morphids appear to be very close to 
the condition apparent when the fundamental dichotomy of the nym- 
phaloids into monocotyledon and dicotyledon feeding groups took 
place. 

Fox (1956: 18-19) considers the Ithomiidae to be as near or 
nearer the Satyridae than to the Danaidae, with which the ithomiids 
had formerly been placed. This conclusion was reached from the 
evidence of various characteristics, particularly of the adults and pu- 
pae. In many respects the ithomiids resemble the Haeterinae of the 
Satyridae, particularly as regards the presence of a vestige of fore- 
wing vein 3 A (a characteristic also shared with the danaids) in both 
groups. The obviously similar tendency of the ithomiids and the 
haeterines to have hyaline wings is, therefore, supported by structural 
evidence. The vestige of forewing vein 3A will serve to separate the 
ithomiids from the remainder of the satyrids, and no discreet andro- 
conial areas are found in the haeterines, but these are usual (and 
taxonomically important) in the ithomiids. All Ithomiidae feed as 
larvae on the dicotyledonous family Solanaceae. The morphids ap- 
pear to be even more closely related to the satyrids than the ithomiids, 
but neither can be combined with the satyrids. 

Family SATYRIDAE Boisduval, 1836 

Satyrides Boisduval, 1836: 166. 
= Hipparchiadae Kirby, 1837: 297. 
= Satyridae Swainson, 1840: 86. 

As mentioned in the preceding section, the Satyridae are related 

MEM. AMER. ENT. SOC, 24 



16 THE SATYRIDAE 

to the nymphaloid families Morphidae and Ithomiidae. 

No single key character will serve to separate the satyrids from 
all other nymphaloids, but a general definition of the Satyridae may 
be formulated as follows: a nymphaloid family having a larva with 
a bifid tail (fleshy anal projections from the eleventh abdominal seg- 
ment) and feeding on monocotyledonous plants and generally having 
adults with the cells of the fore- and hindwings closed by tubular 
veins, forewing veins swollen at their bases and with more or less 
shaggy palpi. Only the habit of feeding on monocots and the bifid 
larval tail are universal in the Satyridae, and some morphids feed on 
monocotyledons. Many other nymphaloids (danaids, ithomiids, ac- 
raeines, etc.) have the cells of both wings closed by tubular veins, 
and the satyrid subfamily Ragadiinae is defined by not having the 
hindwing cell closed by tubular veins, often not closed at all. While 
many of the satyrids have shaggy palpi, many of the primitive ones 
have only very short hairs on the palpi. This part of the definition 
universally applies only to the temperate and arctic species. The 
forewing veins are usually, but not universally, swollen at their bases. 
The bases of the forewing veins of such satyrid genera as Melanitis, 
Lethe, Oeneis and Melanargia are less swollen than are those of such 
"true" nymphalid genera as Mestra, Bolboneura and Callicore (see 
Schatz andRober, 1892). 

The Satyridae are nearly cosmopolitan, being found wherever but- 
terflies in general are found, except on some oceanic islands. These 
butterflies, like most others, find their greatest development and pro- 
liferation in the tropics, not in the temperate zone as claimed by Ehr- 
lich and Raven, (1965: 589). 

Seven subfamilies are recognized in this family. The most primi- 
tive of these are the Haeterinae, Brassolinae and Biinae; intermediate 
are the Elymniinae and Eritinae; the Ragadiinae and Satyrinae are 
advanced. 

In the following pages the individual subfamilies are characterized 
and diagnoses are provided for the tribes embodying the following 
parameters: (1) condition of the eyes; (2) relative length of the an- 
tennae; (3) characteristics of the antennal club; (4) relative lengths 
of the second and third segments of the palpus; (5) the development 
of the hairs of the palpus; (6) the development of the male foreleg; 
(7) the development of the female foreleg; (8) the relative lengths 



LEE D. MILLER 



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18 THE SATYRIDAE 

of the mid- and hindlegs, if significant; (9) the relative lengths of the 
midtibia and the first midtarsal subsegment; (10) the presence or ab- 
sence of dorsal spinosity on the midtibia; (11 ) the development of 
tibial spurs, if any; (12) the presence or absence of a dorsal, distal 
spine at the end of the midtibia; (13) the length and shape of the 
fore wing cell; (14) the branching pattern of the forewing radial veins; 
(15) the relative positions of the origins of torewing veins Rs and 
Mi; (16) the relative positions of the forewing medial veins; (17) 
the relative positions of forewing veins Ms, Cui and Cuy, (18) the 
patterns of inflation of the forewing vein bases; (19) the length and 
shape of the hindwing cell; (20) the relative lengths of hindwing 
veins Sc + Ri and 3 A; (21) the relative positions of hindwing veins 
Mh and Cui; (22) the relative positions of the hindwing medial veins; 
and (23) characteristic and aberrant superficial patterns and the dis- 
tribution of androconia. 

Table 2 gives the character states of certain key characteristics 
within the satyrid subfamilies. A key to these subfamilies follows. 



Key to the Subfamilies of the Satyridae 

1. Hindwing cell open or closed only by aberrantly placed veins (Figs. 134, 

1 39) Ragadiinae 

Hindwing cell closed by tubular veins (Fig. 2, tor example) 2 

2. Forewing with a vestige of vein 3A at base (Figs. 2, 7, 8) Haeterinae 

Forewing vein 3 A fused with 2A throughout its entire length (Fig. 21, for 

example) 3 

3. Hindwing vein R x free basally from Sc + R x (Figs. 9, 90, for example) 4 
Hindwing vein R x completely fused with Sc (Fig. 21, for example) 5 

4. Forewing cell greater than half length of wing; Neotropical species 

Brassolinae 

Forewing cell much less than half length of wing; Paleotropical species .... 
Elymniinae (Elymniini) 

5. Posttarsal claws bifid (Figs. 39. 47) 6 

Posttarsal claws simple 7 

6. Forewing cell more than half length of wing Biinae (Melanitini) 

Forewing cell less than half length of wing 

Elymniinae {Samanta) and Manataria 

7 '. Female foretarsus with fewer than five subsegments many Satyrinae 

Female foretarsus with five subsegments 8 

8. Female foretarsus strongly clubbed distad (Fig. 174, for example) 

many Satyrinae 

Female foretarsus not so clubbed (Figs. 24, 52. 124, for example) 9 



LEE D. MILLER 19 

9. Forewing cell half length of wing or less most Eiymniinae 

Forewing cell greater than half length of wing 10 

10. Forewing cell produced at the origin of M 3 (Figs. 21, 29, for example); 

New World species New World Biinae 

Forewing cell produced at the origin of M 1 (Figs. 121, 127): Indo-Aus- 
tralian species Eritinae 

Subfamily HAETERINAE Herrich-Schaffer, 1864 

Hetaerina [sic] Herrich-Schaffer, 1864: 124. 
= Pierellinae Clark, 1947: 149. 

The Haeterinae on several counts are the most primitive of the 
Satyridae. These butterflies cannot be considered close to any other 
subfamily but show more affinities (albeit slight ones) with the Biinae 
than with the Brassolinae or Eiymniinae. From several aspects — 
the free proximal end of forewing vein 3A, the general elongate wing 
form, the seta-like modification of the wing scales and the preference 
for a deep forest habitat — this subfamily seems to form a connecting 
link with the Ithomiidae (Fox, 1956: 18-19), but, again, this is a 
very remote relationship, if not a completely spurious one arising from 
ecological adaptations. 

In itself the free proximal vestige of forewing vein 3A serves to 
distinguish the haeterines from all other satyrids. This characteristic 
is otherwise shown in the Nymphaloidea by the Ithomiidae, Danaidae, 
some Morphidae (Taenaris) and such nymphalid genera as Colaenis, 
Kallima, Apatwina and the very primitive Callinaga. The general 
shape of the wings (Figs. 2, 7, 8) is characteristic, as is the rounded 
hindwing cell. The combination of a dorsally spinose midtibia and 
the lack of tibial spurs (Fig. 6) is shown by no other satyrid. The 
lack of distal scaling on the antennae is also peculiar to the present 
subfamily. 

All members of the Haeterinae are restricted to the American 
tropics and are found only in the dense forests. 

A single tribe, the Haeterini, includes all members of the sub- 
family. 

The most comprehensive revision of the Haeterinae presently 
available is that of Brown (1942). 

MEM. AMER. ENT. SOC, 24 



20 



THE SATYRIDAE 



Tribe Haeterini Herrich-Schaffer, 1864 
(Figures 1-8) 

Hetaerina [sic] Herrich-Schaffer, 1864: 124. 
= Pierellinae Clark, 1947: 149. 

The diagnostic features of this tribe are those outlined for the 
subfamily above. The Haeterini are characterized as follows: 

The eyes are naked. The antennae vary little in length from 
slightly over one-third (Pierella) to just over two-fifths (Haetera) 
the length of the forewing costa. The antennal club is weakly de- 
veloped, generally less than twice the thickness of the shaft and oc- 
cupying the distal one-fourth to one-fifth of the antenna. The anten- 
nae of the haeterines are scaled on only the proximal few segments: 
the antennae of all other satyrids are fully scaled. The third segment 
of the palpus is usually less than one-fifth the length of the second, 
although the third segment is between one-third and one-fourth the 
length of the second in Haetera and Pseudohaetera. The hairs on 
the second segment of the palpus are generally about as long as the 
segment is wide (the hairs are twice as long as the segment is wide 
in D nice do). 



0.4 x ML, FL 




Haetera 



Cithaerias 



Pierella 



Fig. 1. Haeterinae: Haeterini. Relative lengths of the femur + tibia + 
tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and 
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML 
value is unity. The top bar for each genus represents the measurements ob- 
tained from males, the bottom bar those from females. 



The male foreleg is well developed for a nymphaloid, the tibia is 
longer than the femur, although both are about equal in Pierella, and 
there is a single, unspined tarsal subsegment in all genera. The female 
foreleg is also well developed with a pentamerous, unclubbed tarsus 



LEE D. MILLER 



21 






Figs. 2-8. Haeterinae: Haeterini. 2. Haetera piera (Linne), 6 venation. 
3. H. piera, palpus (in this and all succeeding illustrations of palpi only the 
distal two segments are figured). 4. H. piera, S foreleg (in this and all suc- 
ceeding illustrations of the $ forelegs, unless otherwise indicated, the femur, 
tibia and tarsus are figured. 5. H. piera, 2 foretarsus. 6. H. piera, midleg 
(in this and all succeeding illustrations of the midlegs only the femur, tibia 
and tarsus are figured). 7. Pierella nereis (Drury), $ venation. 8. Cithae- 
rias pireta (Cramer), 6 venation. 

armed with spines on the second, third and fourth subsegments. The 
mid- and hindlegs are of about equal length. The midtibia is one 
and one-half to two times as long as the first midtarsal subsegment, is 
spiny dorsad and lacks tibial spurs or a spine at the dorsal, distal end. 
The relative development of the legs is shown in Fig. 1 . 

The forewing cell varies from just greater than half to more than 
three-fifths the length of the forewing costa and is rounded distad in 
all genera. The forewing radial veins arise in three branches from 
the cell. Forewing veins Rs and Mi are connate or approximate at 



MEM. AMER. ENT. SOC, 24 



22 THE SATYRIDAE 

their origins, vein M2 arises midway between Mi and M3 and vein 
Cui arises midway between M 3 and C112, or slightly nearer the former. 
The forewing vein Sc is inflated in all genera, and the other veins may 
be: both the cubital stem and 2A in Pierella, only 2A in Haetera. 
The presence of a proximal vestige of forewing vein 3A is diagnostic 
in the Haeterinae. 

The hindwing cell is also distally rounded and is from two-fifths 
{Pierella) to three-fifths (Cithaerias) as long as the wing, as measured 
to the origin and end of M3, respectively. Hindwing veins Sc + Ri 
and 3A are of about the same length. The origins of hindwing veins 
M3 and Cui may be widely separated ( Cithaerias, Fig. 8 ) to connate 
on a common stalk (Pierella, Fig. 7 ). Vein M L » arises about midway 
between Mi and M3, or slightly nearer the latter. 

The pattern is distinctive (see Schwanwitsch, 1925 ) : the scales of 
most species are reduced to narrow, hairlike vestiges, resulting in a 
hyaline or translucent appearance in most species. Relatively un- 
modified scales are displayed by some species, particularly of the 
genera Pierella and Haetera, with resultant opaque wings. 



Genera Included in the Haeterini 

Callitaera Butler, 1868b: 101. Type-species: "Papilio aurora Felder" (= Ci- 
thaerias phantoma Fassl, fide Clifton, in litt.), designated by Brown, 
1942: 311. 

Cithaerias Hiibner, [1819] (1816-1826): 53. Type-species: "Papilio andro- 
meda Fabricius" ( = Papilio pireta Cramer, according to Brown, 1942), 
designated by Scudder, 1875a: 143. Papilio menander Drury, often 
regarded as the type-species, was shown by d'Almeida (1951: 496) to 
be identical with Papilio pireta Cramer. 

Dulcedo d'Almeida, 1951: 501. Type-species: Haetera polita Hewitson, by 
original designation. 

Haetera Fabricius, 1807: 284. Type-species: Papilio piera Linne, designated 
by Butler, 1868a: 195. 

= Oreas Hiibner, [1807] (1806-1838): pi. [82]. Type-species: Pa- 
pilio piera Linne, by monotypy. Oreas is sunk to Haetera Fabri- 
cius in accordance with Opinion 137 (1942) of the International 
Commission of Zoological Nomenclature. 

Pierella Herrich-Schaffer, 1865: 65. Type-species: Papilio nereis Drury, desig- 
nated by Butler, 1868a: 195. 

Pseudohaetera Brown, 1942: 330. Type-species: Haetera hypaesia Hewitson, 
by original designation. 



LEE D. MILLER 23 

Subfamily BRASSOLINAE Boisduval, 1836 
Brassolides Boisduval, 1836: 166. 

The brassolines have long been considered a separate family 
somewhat intermediate between the satyrids and the morphids, but 
Ehrlich (1958: 351-352) placed the brassolines with the morphids. 
There is little except the large size of the butterflies to support this 
combination: the larvae of most Neotropical morphids feed on dicots, 
whereas the brassolines feed exclusively on monocots; brassoline lar- 
vae have bifid tails, but the tails are greatly reduced in the morphids. 
These two characters intimately associate the brassolines with the 
other satyrids. The cells of both wings are closed in the brassolines, 
as in the satyrids generally, whereas the hindwing cell is open in most 
of the Morphidae. Within the Satyridae the brassolines are allied to 
the New World Biinae, particularly through such genera as Narope. 
There seems little justification, in view of the above facts, for retain- 
ing the brassolines as a family separate from the Satyridae. 

The brassolines may be distinguished from all other New World 
Satyridae by the basal separation of hindwing veins Sc and Ri (Figs. 
9, 13, 14, 17, 19). This characteristic is also shown by the Old 
World Elymniini (Elymniinae), but the extremely short cell and vir- 
tually connate veins M3 and Cui of the forewing immediately disting- 
uishes the elymniines. The Haeterinae show an approximate condi- 
tion of hindwing veins Sc + Ri and Rs, but there is no basal separation 
of Ri. The brassolines lack the vestige of forewing vein 3 A which 
characterizes the Haeterinae. 

All brassolines are Neotropical, none being found further north 
than southern Mexico. Some species of Caligo are banana pests in 
Central America, and around the banana plantations these insects 
often may be seen flying just at dusk. 

Clark ( 1947, 1948 ) recognized two tribes as subfamilies of "Bras- 
solidae", but no good evidence is found for splitting the brassolines 
into more than a single tribe, the Brassolini. Within the Brassolini 
groupings are possible, but not tribal ones. 

Tribe Brassolini Boisduval, 1836 
(Figures 9-19) 

Brassolides Boisduval, 1836: 166. 
= Caliginae Clark. 1947: 149. 

MEM. AMER. ENT. SOC, 24 



24 



THE SATYRIDAE 




Figs. 9-19. Brassolinae: Brassolini. 9. Brassolis sophorae (Linne), £ 
venation. 10. B. sophorae, palpus. 11. B. sophorae, £ foreleg. 12. B. so- 
phorae, midleg. 13. Opsiphanes cassiae (Linne), £ venation (after Schatz 
and Rober, 1892). 14. Caligo atreus Kollar, £ venation (after Schatz and 
Rober, 1892). 15. C. eurylochus (Cramer), 9 foretarsus. 16. C. eiirylochus, 
midtarsus and distal portion of midtibia. 17. Dasyophthalma rusiiui (Godart), 
£ venation. 18. D. rusina, 9 foretarsus. 19. Narope cyllastros Doubleday 
and Hewitson, £ venation. 

The characteristics of the subfamily are those of the tribe. Four 
groups may be noted within the tribe, but they are only loose as- 
semblages and tend to grade into one another. The diagnostic char- 
acters are summarized in Table 3. 



LEE D. MILLER 



25 



The Brassolis-series, sensu stricto, is composed of robust species 
with naked eyes, moderately well developed or absent tibial spurs and 
a basically brown upper surface with a transverse band of orange, 
yellow or red across the forewing. 

The Caligo-senes includes large, broad-winged, but not robust- 
bodied butterflies with naked eyes, well developed tibial spurs and a 
basically iridescent purple upper surface with faint longitudinal light 
markings. 

The Dasyophthalma-series is comprised of a single genus of broad- 
winged species with hairy eyes, moderately well developed tibial spurs 
and a basically brown upper surface, with or without iridescence, and 
with longitudinal lighter markings. 

The Narope-sevies is also comprised of a single genus of medium- 
sized butterflies with acute forewing costal margins, naked eyes, mod- 
erately well developed tibial spurs and a brown upper surface with 
little or no maculation. 

Table 3 



The character states of four characters ir 


i the genera 


of the Brassolinae 


Brassolini. 








Length of fore- 
wing cell/ 






Antennal 


Tibial 


length of 


Genus 


Eyes 


club 


spurs 


forewing costa 


Brassolis-series, s. str. 










Brassolis 


Naked 


Pronounced 


Present, ex. 


3 5-2/3 








B. sophorae 


Catoblepia 


Naked 


Gradual 


Present 


3 5-2/3 


Dynastor 


Naked 


Gradual 


Present 


1/2-3/5 


Opoptera 


Naked 


Gradual 


Present 


3/5-2/3 


Opsiphanes 


Naked 


Gradual 


Present 


1/2-3 5 


Penetes 


Naked 


Gradual 


Present 


3/5-2/3 


Selenophanes 


Naked 


Gradual 


Present 


3/5-2/3 


Caligo-senes 










Caligo 


Naked 


Gradual 


Present 


3 5-2/3 


Eryphanis 


Naked 


Gradual 


Present 


3/5-2/3 


Dasyophthalma-series 










Dasyophthalma 


Hairy 


Gradual 


Present 


3 5-2/3 


Narope-series 










Narope 


Naked 


Gradual 


Present 


just >l/2 



MEM. AMER. ENT. SOC, 24 



26 THE SATYRIDAE 

The Brassolini are characterized as follows. — The eyes are naked 
in all genera but Dasyophthalma in which they are densely hairy. 
The antennae are short, one-third to two-fifths the length of the fore- 
wing costa, and the antennal club is weakly developed in most genera 
(just over twice the width of the shaft in Brassolis). The third seg- 
ment of the palpus is generally one-fourth to one-sixth the length of 
the second, although it is far shorter in such rather advanced genera 
as Brassolis and Opsiphanes and somewhat longer in the more primi- 
tive Narope. The hairs of the second segment of the palpus are gen- 
erally shorter than the segment is wide. 

The male foreleg is well-developed with a mono- to trimerous tar- 
sus; the femur is longer than the tibia in all genera. The female fore- 
leg is likewise well developed with a pentamerous, unclubbed tarsus 
bearing spines on at least the third and fourth (often the first and 
second, too) subsegments. The midtibia varies from just less than 
twice {Caligo and Eryphanis) to well over twice ( Brassolis-series ) 
as long as the proximal midtarsal subsegment. The midtibia is spiny 
dorsad with well to moderately well developed tibial spurs (except in 
Brassolis sophorae, where the tibial spurs are totally wanting, Fig. 
12); there is no spine at the dorsal, distal end of the midtibia. 

The forewing cell is produced at the origin of Ma, particularly in 
Caligo and Eryphanis. The cell is generally three-fifths to two-thirds 
the length of the forewing costa, although it is somewhat shorter in 
Dynastor, Opsiphanes and Narope. The forewing radial veins arise 
from the cell in three branches. Forewing veins Rs and Mi are well 
separated at their origins; vein M L > usually arises nearer Mi than M3, 
although M2 arises nearer M3 in Caligo and Dynastor; vein Cui arises 
nearer Ma than CU2, except in Dasyophthalma. The forewing veins 
are seldom inflated, though the base of Sc may be in some genera. 

The hindwing cell is produced marginad by a migration of cross- 
vein mriii3 and is generally about three-fifths the length of the wing, 
measured to the origin and the end of Ma, respectively. Hindwing 
veins Sc and Ri are separate proximally in this tribe, as in the Elym- 
niini (Elymniinae); Sc + Ri is longer than 3A. At their origins veins 
Ms and Cui are well separated, and M 3 arises nearer Ma than Mi. 

The upper surface patterns have been described in the analysis of 
the generic series. On the under surface the wings are cryptically 
patterned, light brown with darker striations, much as in the Antir- 



LEE D. MILLER 27 

rhini. In the brassolines, however, there are often large "eye-spots", 
particularly on the hindwing, which have been credited with being 
protective markings (the wings are rapidly opened when the butter- 
fly is threatened by a predator, giving the impression of large eyes 
blinking). The development of these eye-spots gives the common 
name of the Cali go-series, the "owl butterflies". 

Genera Included in the Brassolini 
The Brassolis-series, sensu strict o 

Brassolis Fabricius, 1807: 282. Type-species: Papilio sophorae Linne, desig- 
nated by Westwood, 1851, in Doubleday, Westwood and Hewitson, 
1846-1852: 341. 

Catoblepia Stichel, 1902: 488, 491. Type-species: Brassolis amphiroe Hiibner, 
by original designation. 

Dynastor Westwood and Hewitson, 1849, in Doubleday, Westwood and Hewit- 
son, 1846-1852: pi. 58. Type-species: Dynastor napoleon Westwood 
and Hewitson, by monotypy. 

Opoptera Aurivillius, 1882: 75. Type-species: Brassolis syme Hiibner, by origi- 
nal designation. 

Opsiphanes Westwood and Hewitson, 1849, in Doubleday, Westwood and Hew- 
itson, 1846-1852: pi. 57. Type-species: Opsiphanes sallei Westwood 
and Hewitson, designated by Scudder, 1875a: 233. 

Penetes Westwood and Hewitson, 1849, in Doubleday, Westwood and Hewit- 
son, 1846-1852: pi. 58. Type-species: Penetes pamphanis Westwood 
and Hewitson, by monotypy. 

Selenophanes Staudinger, 1888: 212. Type-species: Papilio cassiope Cramer, 
by original designation. 

The Caligo-sehes 

Caligo Hiibner. [1819] (1816-1826): 51. Type-species: Caligo eurylochus 

Hiibner, designated by Scudder, 1875a: 129. 
Eryphanis Boisduval, 1870: 57. Type-species: Papilio automedon Cramer, by 

monotypy. 

Dasyophthalma-series 

Dasyophthalma Westwood, 1851, in Doubleday, Westwood and Hewitson, 
1846-1852: 343. Type-species: Brassolis rusina Godart, designated by 
Scudder, 1875a: 155. 

MEM. AMER. ENT. SOC, 24 



28 THE SATYRIDAE 

/Varo/^-series 

Narope Westwood and Hewitson, 1 849, in Doubleday, Westwood and Hewit- 
son, 1846-1852: pi. 50. Type-species: Narope cyllastros Westwood 
and Hewitson, by monotypy. 

Subfamily BIINAE Herrich-Schaffer, 1864 
Biina Herrich-Schaffer, 1864: 124. 

The Biinae are the third of the very primitive satyrid groups and 
show no especially close relationship with any other subfamily. The 
Neotropical biines share some vague affinities with the haeterines and 
some firmer ones with the brassolines, particularly as regards vena- 
tion, whereas the Paleotropical biines grade toward the Lethini of the 
next subfamily. 

The biines are generally large to very large satyrids, often brightly 
colored above and always cryptically patterned beneath with few and 
usually poorly developed eye-spots on either wing surface. The long 
forewing cell, generally three-fifths the length of the wing except in 
some melanitines (Fig. 34), is characteristic, and the well separated 
hindwing veins M3 and Cui serve to distinguish these butterflies from 
most elymniines. The wings are never translucent or transparent, as 
in the haeterines. 

Table 4 
Some of the diagnostic characters of the tribes of the Biinae. 

Character Antirrhini Biini Melanitini 

Tarsal claws Simple Simple Bifid 

Tibial spurs Present, but Absent Present 

poorly devel- 
oped 
Length of 3rd seg. of 
palpus relative to 

length of second <l/4 >l/2 1/4 

Forewing radial veins 3 branches 1 branch, 3 branches 

aberrant 

Inflation of forewing veins .. Only Sc, if any; .. Sc, Cu, 2A Only Sc, slight 

or else v. slight greatly inflated 



LEE D. MILLER 29 

All biines are tropical. The Antirrhini and Biini are found ex- 
clusively in the Neotropics and are forest dwellers, whereas the Mel- 
anitini are Paleotropical (from West Africa to Australia) and are 
found in less dense situations, though many of these species are cre- 
puscular. 

The Biinae are split, somewhat reluctantly, into three tribes, but 
Bia is far too aberrant to be referred to either of the other two. The 
diagnostic features of the three tribes are given in Table 4, and a key 
to these tribes is given below. 

Key to the Tribes of the Biinae 

1. Tarsal claws bifid (Fig. 39); Paleotropical species Melanitini 

Tarsal claws simple; Neotropical species 2 

2. Tibial spurs present, though poorly developed (Fig. 25); third segment of 

palpus less than one-fourth length of second segment (Fig. 22) 

Antirrhini 

Tibial spurs absent (Fig. 33); third segment of palpus greater than half 
length of second segment (Fig. 30) Biini 

The New World members of the Biinae are considered to be more 
primitive because of ( 1 ) their greater convergence toward the Bras- 
solinae and Haeterinae, (2) the greater relative size of the male fore- 
leg and (3) the unclubbed female foreleg. The Melanitini are the 
more advanced members of the subfamily and seem to show relation- 
ships with the more advanced Elymniinae. 

Tribe Antirrhini, new tribe 
(Figures 20-28) 

The simple tarsal claws serve to separate this New World tribe 
from the Old World Melanitini. Many characteristics serve to sep- 
arate the Antirrhini from the Biini, notably the presence of tibial 
spurs, the more reduced third segment of the palpus, the arising of 
the forewing radial veins in three branches and the relative placement 
of several other veins on the cells. 

The Antirrhini are Neotropical insects, found from Guatemala to 
Paraguay and northern Argentina. All seem to be forest species, 
seldom venturing into open areas. The tribe Antirrhini is character- 
ized as follows: 

The eyes are naked in all genera but Sinarista, in which they are 
sparsely hairy. The antennae are about half as long as the forewing 

MEM. AMER. ENT. SOC, 24 



30 



THE SATYRIDAE 



0.4xML, 



FL 



— ■+ 



l_ 



x; 



'///////> 



ML 

ANTIRRHINI 



HL 



BIINI 




IxML 



Antirrhea 



Caerois 



Triteleuta 



Bia 



Melanitis 



Hipio 



Cyllogenes 



Parantirrhoea 



Fig. 20. Biinae: Antirrhini, Biini and Melanitini. Relative lengths of the 
femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light 
stippling) and hindlegs (HL, heavy stippling) of selected genera. In all in- 
stances the ML value is unity. The top bar for each genus represents the mea- 
surements obtained from males, the bottom bar those from females. 



costa, and the antennal club is weakly developed (it is completely in- 
distinct in Triteleuta). When the club is distinct it occupies the dis- 
tal one-third to one-quarter of the antenna and is seldom more than 
twice as thick as the shaft. The third segment of the palpus is one- 
fourth to one-sixth the length of the second segment, and the hairs of 
the second segment are about as long as the segment is wide, although 
the hairs of Sinarista are about twice as long as the width of the seg- 
ment. 

The male foreleg is well developed with a monomerous, unspined 
tarsus, and the tibia is usually slightly longer than the femur, although 
the femur is much longer in Caerois. The female foreleg is also well 
developed with a pentamerous, unclubbed tarsus bearing spines on 
the first four subsegments. The midtibia is more than twice as long 



LEE D. MILLER 31 

as the basal midtarsal subsegment, smooth dorsally and bearing weakly 
developed tibial spurs. There is no spine at the dorsal, distal end of 
the midtibia. The relative lengths of the legs are shown in Fig. 20. 

The forewing cell is square-cut in Caerois and Sinarista, but deeply 
excavate at Mi in the other genera. The cell is about three-fifths the 
length of the forewing costa. The radial veins arise in three branches 
from the forewing cell, and veins Rs and Mi are connate to approxi- 
mate at their origins. Forewing vein M- arises midway between Mi 
and M ;i , whereas Cu t usually arises about midway between M:t and 
CU2 (nearer M3 in Caerois). The forewing veins are only slightly 
inflated, if at all; any thickening is at the base of Sc. 

The hindwing cell is highly variable, from very short and narrow 
in Antirrhea to over three-fifths the length of the wing, measured to 
the end of M3, and rounded in Caerois. Hindwing vein Sc + Ri is 
usually longer than 3A, although the two veins are of about the same 
length in Sinarista. Hindwing veins M3 and Cui are generally well 
separated at their origins, but they are approximate in Antirrhea; vein 
M2 arises midway between Mi and M3. 

Most of the species are rather brightly colored above with few 
ocelli, although some species, such as Sinarista adopt iva Weymer, 
have rows of submarginal ocelli. On the under surface all species 
are cryptically marked with many fine, dark striations on a tan to 
brown ground color. There are androconial hair patches on the under 
surface of the forewing along vein 2 A in all genera but Sinarista, and 
these androconial areas frequently result in distortions of the venation. 

Genera Included in the Antirrhini 

Antirrhea Hiibner, [1822] (1806-1838): pi. 294. Type-species: Antirrhea ar- 
chaea Hiibner, by monotypy. 

= A nchiphlebia Butler, 1868b: 106. Type-species: Antirrhea archaea 
Hiibner, by original designation. 
Caerois Hiibner, [1819] (1816-1826): 56. Type-species: Papilio arcesilaus 
Cramer (= Papilio chorinaeus Fabricius). by monotypy. 

= Arpidea Duncan, 1837: 180. Type-species: Papilio chorinaeus 
Fabricius, by monotypy. 
Sinarista Weymer, 1909: 164. Type-species: Sinarista adoptiva Weymer, by 

monotypy. 
Triteleuta Strand, 1912: 44. Type-species: Antirrhea tomasia Butler, by origi- 
nal designation. Proposed as a subgenus of Antirrhea Hiibner. 

MEM. AMER. ENT. SOC, 24 



32 



THE SATYRIDAE 







Figs. 21-28. Biinae: Antirrhini. 21. Antirrhea archaea Hiibner, $ vena- 
tion. 22. A. archaea, palpus. 23. A. archaea, $ foreleg. 24. A. archaea, 9 
foretarsus. 25. A. archaea, midleg. 26. Caerois chorinaeus (Fabricius), $ 
venation. 27. C. chorinaeus, $ foreleg. 28. Sinarista adoptiva Weymer, $ 
venation. 

Tribe Biini Herrich-Schaffer, 1864 
(Figures 20, 29-33) 

Biina Herrich-Schaffer, 1864: 124. 

Bia actoriaena (Linne), the only species included in the Biini, is 
the most aberrant member of the Satyridae. It has been considered 
variously a brassoline (Weymer, 1912; Clark, 1947, 1948) or a per- 
fectly good satyrid (Rober, 1892; Ehrlich, 1958). Bia is definitely 



LEE D. MILLER 33 

a satyrid and shows relationships to other members of the Biinae, 
chiefly through pattern convergence, but it is far too aberrant to be 
included in either of the other two biine tribes. This species can be 
distinguished from all other biines by the lack of tibial spurs, in addi- 
tion to a multitude of other characteristics enumerated below in the 
description of the tribe. Bia is primitive, though highly specialized, 
and may be characterized as follows: 

The eyes are naked. The antennae are half as long as the fore- 
wing costa, and the club is poorly developed occupying the distal 
quarter of the antenna and about twice as thick as the shaft. The 
third segment of the palpus is very long, over half the length of the 
second segment, and the hairs of the second segment are not as long 
as the segment is wide. 

The male foreleg is well developed, with the femur longer than the 
tibia and a monomerous, unspined tarsus. The female foreleg is also 
well developed with a pentamerous, unclubbed tarsus bearing spines 
on the first four subsegments. The midtibia is smooth dorsally, more 
than twice as long as the proximal midtarsal subsegment and armed 
with neither tibial spurs nor a spine at the dorsal, distal end. 

The forewing cell, measured to the origin of M3, is greater than 
three-fifths the length of the forewing costa, but the cell is deeply ex- 
cavate at the origins of Mi and M2, and the cell is only about half 
the length of the forewing costa when it is measured to the origin of 
Mi. All the forewing radial veins arise from a single branch from 
the cell, and Ri anastomoses distally with Sc, resulting in a confused 
venational pattern along the costa. Forewing veins Rs and Mi are 
well separated at their origins. Vein M2 arises much nearer Mi than 
M s , whereas Cui arises midway between M3 and Cui>. Veins Sc, the 
cubital stem and 2A are all strongly inflated at their bases, the in- 
flation of the latter suggesting the pattern shown in the Mycalesini of 
the next subfamily. 

The hindwing cell is strongly produced at the origin of Ms and 
about three-fifths the length of the wing measured to the end of M3. 
Hindwing vein Sc + Ri is longer than 3 A. Veins M3 and Cui are 
well separated at their origins, and Ml> arises nearer Mi than M3. 

The upper surface is dark brown with patches of orange and iri- 
descent blue scales. On the under surface Bia is marked like other 
members of the subfamily: tan with numerous, longitudinal brown 

MEM. AMER. ENT. SOC, 24 



34 



THE SATYRIDAE 







Figs. 29-33. Biinae: Biini, all Bia actoriaena (Linne). 
30. palpus. 31. A foreleg. 32. 2 foretarsus. 33. midleg. 



29. £ venation. 



striae. The androconial areas on the hindwing are characteristic: 
there is a patch of mealy scales on the upper end of the cell along 
crossvein r s -nii, and a long hair tuft lies along 2A. 

Genus Included in the Biini 

Bia Hubner, [1819] (1816-1826): 51. Type-species: Papilio actoriaena Linne, 
by monotypy. 

Tribe Melanitini, new tribe 
(Figures 20, 34-40) 

The melanitines are the most advanced members of the Biinae, as 
noted above, particularly with regard to the more reduced male fore- 
leg and the slightly clubbed female foretarsus (Fig. 37). There are, 
in addition to the already-mentioned affinities to the Antirrhini, some 
rather suggestive ties between the Melanitini and the Lethini of the 
next subfamily, especially through the genera Parantirrhoea (Mela- 
nitini) and Ptychandra and Samanta (Lethini). The venation of 
Pty chandra and its pattern relate rather closely (but with modifica- 
tions) to those of the Melanitini, and Samanta displays the bifid claws 
characteristic of the melanitines. Bifid claws are the most character- 
istic feature of this tribe and serve to distinguish it from all other 
satyrids except Samanta and Manataria (a Neotropical genus of un- 
certain position). 



LEE D. MILLER 



35 



All members of this tribe are Paleotropical, being found from 
West Africa through the Malay Peninsula east as far as New Guinea 
and northern Australia. At least some of the species are crepuscular, 
and they do not appear to be so limited to the deep forest as are the 
Neotropical Antirrhini. The Melanitini are characterized as follows: 

The eyes are naked. The antennae are rather uniformly just over 
two-fifths the length of the forewing costa, and the antennal club is 
weakly developed, occupies the distal quarter of the antenna and is 
thickened twice to two and a half times the thickness of the shaft. 
The third segment of the palpus is about one-fourth the length of the 
second, and the hairs of the second segment are much shorter than 
the segment is wide. 




Figs. 34-40. Biinae: Melanitini. 34. Melanitis leda (Linne), 6 venation. 
35. M. leda, palpus. 36. M. leda, S foreleg. 37. M. leda, 2 foretarsus. 38. 
M. leda, midleg. 39. M. leda, bifid tarsal claw. 40. Hipio constantia (Cramer), 
venation of proximal portion of 6 wings. 



The male foreleg is well developed with the femur and the tibia 
of about the same length (the femur is longer in Cyllogenes and 
Parantirrhoea) and bearing a single, usually unspined (weakly so in 
Cyllogenes) tarsal subsegment. The female foreleg is also well de- 
veloped with a pentamerous, unclubbed or weakly clubbed tarsus. 
The fourth tarsal subsegment always bears spines, and the spines are 

MEM. AMER. ENT. SOC, 24 



36 THE SATYRIDAE 

present on the third and fourth subsegments in Gnophodes and Par- 
antirrhoea, on the second, third and fourth in Melanitis and Hipio. 
The midtibia is greater than twice as long as the proximal midtarsal 
subsegment, is smooth dorsad and bears moderately well developed 
tibial spurs but no spine at the dorsal, distal end. Relative lengths of 
legs are shown in Fig. 20. 

The forewing cell varies from just over half to nearly three-fifths 
as long as the forewing costa and is somewhat excavate at ni2-m3. 
The forewing radial veins arise in three branches. Veins Rs and Mi 
are usually well separated at their origins, vein M- arises much closer 
to Mi than to M3 and Cui usually arises midway between M:s and Cui;, 
although Cui arises noticeably nearer Cu^ in Melanitis. The forewing 
veins are little inflated; any inflation is restricted to the base of Sc. 

The hindwing cell is more or less rounded distad and is two-fifths 
to half the length of the wing measured to the end of M3. The hind- 
wing vein Sc + Ri is longer than 3A. Hindwing vein M2 arises mid- 
way between Mi and M3, and veins M3 and Cui are well separated at 
their origins. 

The pattern is similar to that of the Antirrhini with a shaded 
brown and orange upper surface and cryptic coloration on the under 
surface with a brown ground color and fine, dark striations. 

Genera Included in the Melanitini 

Cyllogenes Butler, 1868b: 6. Type-species: Melanitis suradeva Moore, by 
monotypy. 

Gnophodes Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 
1852: 363. Type-species: Gnophodes parmeno Westwood, designated 
by Butler, 1868a: 194. 

Hipio Hubner, [1819] ( 1816-1826): 56. Type-species: Papilio constantia Cra- 
mer, designated by Butler, 1867a: 279. 

Melanitis Fabricius, 1807: 282. Type-species: Papilio leda Linne, designated 
by Butler, 1868a: 194. 

= Cyllo Boisduval, 1832: 140. Type-species: Papilio leda Linne, 
designated by Scudder, 1875a: 151. 

Parantirrhoea Wood-Mason, 1880: 248. Type-species: Parantirrhoea mar- 
shall ii Wood- Mason, by monotypy. 

Genus MANATARIA Kirby, 1908 

(Figures 41-47) 

Manataria Kirby, 1908: 57. Type-species: Tisiphone hercyna Hubner, by 
monotypy. 



LEE D. MILLER 



37 



0.4 x ML, FL 


ML 


HL 


IxML 


W//////A 




^MMMM:- ssHxSg 


V//////////A 






■:■:■:■:-:■:■:■:■:':■:■:::-::-:-:■:•:-:-:■;■;] 


I 


41 







Manataria 




Figs. 41-47. Manataria hercyna (Hiibner). 41. Relative lengths of the 
femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light 
stippling) and hindlegs (HL, heavy stippling). The ML value is unity. The 
top bar represents the measurements obtained from the male, the bottom bar 
those from the female. 42. $ venation. 43. palpus. 44. £ foreleg. 45. 9 
foretarsus. 46. midleg. 47. bifid tarsal claw. 



This South American genus has some morphological and pattern 
features which connect it to the Neorina-series of the Lethini. If this 
were its position, Manataria would be the only Neotropical lethine 
and not particularly closely related to the few Nearctic members of 
the tribe. The bifid tarsal claws, on the other hand, connect the pres- 
ent genus with the sympatric Antirrhini (although this is through the 
Paleotropical Melanitini), but in most other respects Manataria has 
little in common with the antirrhines. The pattern is quite typical of 
the Neorina-series, even to the basad displacement of the ocellus in 
hindwing space Rs-Mi of the under surface. Were it not for the vast 
geographical isolation and the bifid tarsal claws I would place Mana- 
taria in the Lethini, perhaps in a monotypic series, but it seems more 
advisable to call attention to this aberrant genus here without assign- 
ing it to a tribe, pending further information. Manataria is char- 
acterized as follows: 

MEM. AMER. ENT. SOC, 24 



38 THE SATYRIDAE 

The eyes are naked. The antennae are about half the length of 
the forewing costa; the antennal club is poorly developed and less 
than twice as thick as the shaft. The third segment of the palpus is 
just under one-third the length of the second. The hairs of the second 
palpal segment are not as long as the segment is wide. 

The male foreleg is well developed with the tibia slightly longer 
than the femur and a monomerous, unspined tarsus. The female 
foreleg is also not significantly miniaturized with a pentamerous, un- 
clubbed tarsus which is spined on the second, third and fourth sub- 
segments. The midleg is slightly longer than the hindleg (Fig. 41). 
The midtibia is just less than twice as long as the proximal midtarsal 
subsegment, is slightly spinose dorsad, bears no spine at the dorsal, 
distal end and has well developed tibial spurs. The posttarsal claws 
are bifid. The relative lengths of the legs are shown in Fig. 41. 

The forewing cell is produced at the origin of Ms and about half 
as long as the forewing costa. The forewing radial veins arise in 
three branches from the cell, and Rs and Mi are connate. Vein M2 
arises nearer Mi than M3, and Cut arises slightly nearer M3 than CU2. 
Forewing veins Sc and the cubital stem are slightly inflated; 2A is not. 

The hindwing cell is produced at the origin of Ms and is slightly 
more than two-fifths the length of the wing measured to the end of 
M3. Veins Sc + Ri and 3A are of about the same length, and M 3 
and Cui are separate, but approximate, at their origins. Vein M 2 
arises much nearer Mi than Ms. 

The upper surface is dark brown with a transverse yellow spot- 
band on the forewing outside the cell. Below the pattern of the upper 
surface is repeated; in addition, there are pale scrawlings at the apex 
of the forewing and all over the hindwing and ocelli are developed in 
space M1-M2 of the forewing and in all spaces from Rs-Mi to Cu 2 -2A 
of the hindwing. The ocellus in hindwing space Rs-Mi is displaced 
basad as in most of the Lethini. 

Subfamily ELYMNI1NAE Herrich-Schaffer, 1864 

Eurytelidae Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 

1852: 403 (in part). 
Elymniina Herrich-Schaffer, 1864: 124. 
= Enodiinae Clark, 1947: 149. 
= Lethinae Clark, 1948: 77. 



LEE D. MILLER 39 

The Elymniinae are the now dominant primitive satyrid subfamily 
and apparently provided the source for the Satyrinae, Eritinae and 
probably the Ragadiinae — although the origin of the latter subfamily 
is still a problem. In some respects the elymniines may be connected 
to, and may have arisen from, the Melanitini (Biinae) through such 
genera as Ptychandra and Samanta (Lethini), as discussed under the 
melanitines. 

Although not all Elymniinae show this characteristic and some 
that are not elymniines do, most satyrids with the hindwing veins M3 
and Cui connate or closely approximate at their origins are probably 
referable to this subfamily. The form of the hindwing cell is also 
distinctive: it is usually greatly produced marginad at the origin of 
vein M3, although this is not true of some Elymniini because of dis- 
placement of the venation by the androconial patch. The presence of 
androconial patches in specific locations is characteristic of the tribes. 
With the exception of a few lethine genera of the Pararge-series, the 
forewing cell is never more than half the length of the wing, in marked 
contrast with the preceding three subfamilies where the cell is seldom 

Table 5 
Some of the diagnostic characters of the tribes of the Elymniinae. 

Character Lethini Zetherini Elymniini Mycalesini 

Tibial spurs .. Present Present Absent Present 

Eyes Hairy, except ..Naked Naked Naked or hairy 

in Neorina- 
series 
Forewing 

vein 2A .... Little, if at Not inflated .... Not inflated .... Subquadrate 

all, inflated basal infla- 
tion, except 
two genera 
Forewing me- 
dian veins .. M 2 arises M L > arises much M-, arises much IVL arises much 

nearer M x nearer M^_ nearer M 1 nearer M t 

than M 3 than M 3 than M 3 than M 3 

Forewing cell Rounded, ca. .. Slightly exca- .. Strongly exca- .. Strongly exca- 
1 2 length vate, <1 2 vate, <1 2 vate, ca. 1/2 

of forewing length of length of length of 

forewing forewing forewing 

MEM. AMER. ENT. SOC, 24 



40 THE SATYRIDAE 

much less than three-fifths the length of the forewing costa. The male 
foreleg is moderately well developed in all genera and is characteristi- 
cally tapered to a rather sharp point. 

The metropolis of the Elymniinae is the Paleotropical area, and 
three of the four tribes are found more or less exclusively there. The 
other tribe, the Lethini, reaches its greatest development in the Indo- 
Malayan region but has representatives in Africa, the Palearctic and 
two genera in North America. 

The tribes of the Elymniinae all more or less grade into one an- 
other. They are the Lethini, Zetherini, Elymniini and Mycalesini. 
It is difficult to determine which of the tribes is primitive and which 
is derived, if any are, but the A eropetes-series of the Lethini are defin- 
itely primitive, whatever the condition of the other members of the 
tribe; the other tribes have about equal status. A key to the tribes 
of the Elymniinae is given below and an analysis of them in Table 5. 

Key to the Tribes of the Elvmniinae 

1. Tibial spurs absent (Fig. 96) Elymniini 

Tibial spurs present (Fig. 53, for example) 2 

2. Forewing vein 2A inflated at base to a subquadrate thickening (Figs. 99, 

115, for example) most Mycalesini 

Forewing vein 2A, if thickened, not subquadrate at base 3 

3. Eyes hairy; forewing cell frequently rounded distad (Fig. 49. for example) 

Lethini (except Neoriiia-series ) 

Eyes naked; forewing cell frequently excavate (Fig. 90, for example) .. 4 

4. Base of forewing vein Sc about three times the thickness of any other 

stalk (Fig. 106) Mycalesini (Orsotriaena, Bletogona) 

Bases of forewing veins little inflated, Sc seldom over twice the thickness 
of any other vein 5 

5. Forewing vein M., arising much nearer M x than M 3 (mj-m. one-fourth or 

less as long as m 2 -m 3 ; Figs. 81, 86, 87) Zetherini 

Forewing arising nearer, but not much nearer, M x than M 3 

Lethini (Neorina group) 

Tribe Lethini Clark, 1948 
(Figures 48-80) 

Enodiinae Clark, 1947: 149. 
Lethinae Clark, 1948: 77. 

"Lethinae" is used in preference to the older "Enodiinae" for a 
variety of reasons. Enodia is generally considered at best a subgenus 



LEE D. MILLER 41 

of Lethe, but this is an invalid reason for the suppression of a family- 
group name, according to Art. 40 of the International Code of Zoo- 
logical Nomenclature (1961). Part (a), however, of the same Ar- 
ticle states, "If a family-group name, changed before 1961 because 
of such synonymy, has won general acceptance, it is to be maintained 
in the interests of stability." This is precisely applicable to the ques- 
tion at hand. Dos Passos (1964: 99) uses the collective name 
"Lethinae" in preference to the older "Enodiinae". To now replace 
Lethinae (or in this instance, "Lethini") with Enodiinae (Enodiini) 
would not serve the interests of stability of nomenclature. Lethe is 
further to be preferred, all other things being equal, inasmuch as this 
genus is recognized throughout much of the world; Enodia, where it 
is even recognized today, is known only in the eastern part of North 
America. 

The Lethini may be the focal point of much of the evolution of 
the higher categories of the Satyridae. As mentioned earlier in the 
discussion of the Melanitini, there are some vaguely but intriguingly 
indicated connections between that group and the lethines through 
such lethine genera as Ptychandra and Samanta. The venation of 
the former genus, while highly modified by androconial displacement 
of the forewing veins, is suggestive of the pattern in the melanitines 
(Fig. 57). Samanta, while a perfectly typical lethine from almost 
all other aspects, does show the bifid tarsal claws which characterize 
the Melanitini and the Neotropical aberrant genus Manataria. With- 
in the Elymniinae, too, some interesting gradations may be seen. The 
line separating the Zetherini from the Lethini is a thin one, based 
chiefly on the relative positions of some veins and naked versus hairy 
eyes, respectively. The zetherines, in turn, have many characteristics 
in common with the Elymniini, as will be shown below. The genus 
Mandarinia is a puzzle: the distribution of androconia suggest that it 
is a mycalesine, but the lack of inflation of the forewing vein 2A and 
the distally rounded forewing cell are lethine characteristics: this genus 
is provisionally placed in the Lethini (Fig. 73). The African mem- 
bers of the A eropetes-series show yet another relationship. If one 
considers these butterflies and members of the satyrine tribe Dirini 
together, one can almost obtain a graded series from perfectly good 
Lethini to typical Satyrinae. This is probably convergence, as will 
be shown later. 

MEM. AMER. ENT. SOC, 24 



42 THE SATYRIDAE 

The Lethini are generally characterized by the hairy eyes which 
separate them from the Elymniini and the zetherines, the presence of 
tibial spurs (a characteristic of the Elymniini is the lack of tibial 
spurs) and by the more or less rounded forewing cell, never exca- 
vate as in the elymniines and most mycalesines. Most Lethini show 
a characteristic basad displacement (diastoma) of the hindwing ocel- 
lus in space M1-M2. There are a number of generic series, however, 
so it is difficult to distinguish all lethines by a single set of criteria. 

The A eropetes-series, consisting of two genera, is confined to 
South Africa and characterized by the large size, extremely primitive 
pattern (Schwanwitsch, 1924), hairy eyes, connate hindwing veins 
Mi and Cui, extremely spiny midtibia and lack of androconial patches 
(Figs. 67-72). 

The Lethe-series, sensu stricto, has twenty-five nomenclatorially, 
though not necessarily biologically, valid genera, twenty-two of which 
are Indo-Australian or sub-Palearctic, one East African and two North 
American. These insects are medium to large-sized, have a rather 
primitive pattern on the under surface, though not necessarily on the 
upper, hairy eyes, connate to separate hindwing veins M3 and Cui, 
dorsally smooth to only slightly spiny midtibia and frequently with 
androconial patches in the usual positions (Figs. 49-59, 66). 

The Pararge-series has nine Palearctic genera which differ from 
the Lethe-series chiefly in the length of the forewing cell: in all other 
groups the cell is about half the length of the wing, whereas in Par- 
arge and its allies it is about three-fifths as long ( Figs. 60-65 ) . 

The Mandarinia-series is monotypic, and is characterized by its 
Mycalesis-iike androconial area and is found exclusively in China 
(Figs. 73-76). 

The N eorina-series is composed of four Indo-Australian genera. 
These butterflies are characterized by their large size, highly modified 
pattern, naked eyes, spiny midtibia and lack of androconia (Figs. 
77-80). 

The Lethini are characterized as follows: 

The eyes are hairy in all members except those of the Neorina- 
series, which have naked eyes. The antennae are generally from two- 
fifths to nearly half as long as the forewing costa, but the antennae 
are slightly longer than half the costa in such genera as Hermias, 
Tansima and Pararge. The antennal club is moderately well devel- 



LEE D. MILLER 



43 



0.4xML, FL 



I J 



[ 



__ 



__ 



»» 



IZI 






r 



V////A 









,A 



J 



*m 



ML | HL 

Aeropetes series 






Lethe 'series, s. s. 



xML, 



i 






*Sj : 



w 



Z? 




Pararge 



i 

_........:..:..^_. ; _._.w ..i~. — J 



h 



..._j 



Mandarinia I series 



^^ 



z zz 



e^% 



:.,.i 



. ..■.■.■■.■ ■ ■ ■. ■■ ■■ ■ ■ ■■■ '.'.■.■.;.' ■'. 



: : : : : :v:': : : : : : :':^ 



Neorina series 

k : : : : :j : : : : : : : : : : 



Aeropetes 
Paralethe 

Lethe 

Aphysoneura 

Satyrodes 

Enodia 

Zophoessa 

Ptychandra 

Pararge 

Orinoma 

Kirinia 

Mandarinia 

Neorina 
Anadebis 



Fig. 48. Elymniinae: Lethini. Relative lengths of the femur + tibia + 
tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and 
hindlegs (HL. heavy stippling) of selected genera. In all instances the ML 
value is unity. The top bar for each genus represents the measurements ob- 
tained from males, the bottom bar those from females. 



MEM. AMER. ENT. SOC, 24 



44 THE SATYRIDAE 

oped, occupying the distal quarter of the antenna and inflated two to 
three times the thickness of the shaft. The relative lengths of the 
second and third segments of the palpus are highly variable within 
the tribe: the third segment is less than one-sixth the length of the 
second in Crebeta and between one-third and one-half in Hermias; in 
most genera the third palpal segment is one-fourth to one-fifth the 
length of the second. The length of the hairs of the second segment 
of the palpus is also variable: the hairs are shorter than the segment 
is wide in Hermias, but four times as long as the segment is wide in 
such genera as Magula and Lopinga; in most genera the hairs are 
about three times as long as the second segment is wide. 

The development of the male foreleg is also variable, although 
the leg itself is relatively unminiaturized. There is a great variation 
in the relative length of the forefemur and tibia, and the tarsus has 
one to three subsegments and is usually unspined, although there are 
many tarsal spines in such genera as Ethope and Paralethe. The 
female foreleg is likewise well developed and slightly clubbed at the 
end of the tarsus. The tarsus is always pentamerous with spines on 
at least the second, third and fourth subsegments (also on the first 
subsegment in such genera as Lethe and Ptychandra). The mid- 
and hindlegs are of about the same length. The midtibia is always 
less than twice the length of the first midtarsal subsegment. In the 
South African Aeropetes and Paralethe the midtibia is extremely 
spinose dorsad (Fig. 71 ), but the midtibia is smooth in other genera, 
such as Lethe and Satyrodes (Fig. 53). The tibial spurs are always 
present and well developed, but there is no spine at the dorsal, distal 
end of the midtibia. The relative lengths of the legs are shown in 
Fig. 48. 

The following cell is square-cut or rounded distad, never excavate 
as in other elymniine tribes, and about half as long as the forewing 
costa (Fig. 49, for example) — but this characteristic is by no means 
universal. In Ptychandra the cell is only a third the length of the 
wing (Fig. 57), and in most members of the Pararge-series the cell 
is about three-fifths the length of the costa ( Figs. 60, 64, 65 ) . The 
forewing radial veins always arise in three branches from the cell, and 
veins Rs and Mi are usually well separated at their origins, although 
they may arise approximate to one another. Forewing vein M2 arises 
nearer, though not much nearer, Mi than M 3 ; and Cui arises midway 



LEE D. MILLER 



45 







Figs. 49-56. Elymniinae: Lethini. 49. Lethe europa (Fabricius), S vena- 
tion. 50. L. europa, palpus. 51. L. europa, 6 foreleg. 52. L. europa, 9 fore- 
tarsus. 53. L. europa, midleg. 54. Raugbia scanda (Moore), 6 venation. 
55. Rhaphicera satricus (Westwood and Hewitson), £ venation. 56. Tansima 
satyrina (Butler), £ venation. 

between Mb and C112 in most genera, but Cui arises nearer Cu- in 
several genera (e.g., Lethe, Fig. 49, Ptychandra, Fig. 57, and Pararge, 
Fig. 60). Forewing vein Sc is often somewhat inflated, the cubital 
stem seldom inflated, although it is much inflated in Ptychandra, and 
2 A is never much swollen. 

The hindwing cell is produced at the origin of vein Ms and is 



MEM. AMER. ENT. SOC, 24 



46 



THE SATYRIDAE 



two-fifths to three-fifths the length of the wing measured to the end of 
Ma. Hindwing vein Sc + Ri is of equal length or shorter than 3 A. 
Veins M3 and Cui are usually connate, but they are well separated 
in such aberrant genera as Mandarinia, Ptychandra and Rhaphicera. 
Vein Mi usually arises nearer Mi than Ms, though not much nearer. 
In Aeropetes M 2 arises midway between Mi and M3, slightly nearer 
Ma in the Pararge-sehes and much nearer Ms in Mandarinia. 

The pattern is highly variable, the main unifying feature being the 
diastoma of the hindwing ocelli in space M1-M2 which was mentioned 






Figs. 57-59. Elymniinae: Lethini. 57. Ptychandra lorquinii Felder and 
Felder, £ venation. 58. Enodia portlandia (Fabricins), 6 venation. 59. Saty- 
rodes eurydice (Linne), $ venation. 

earlier. This characteristic is by no means universal, though, and is 
lacking in many genera, either because of the loss of many ocelli or 
extensive realignment of all the hindwing spots. There are several 
sites of androconial generation in the Lethini. Forewing patches are 
uncommon and restricted to the area between the origin of the cubitus 
and the inner margin. These forewing patches are rarely hair tufts 
(Ptychandra, Fig. 57) and are more frequently small patches of mealy 
scales under the origin of Cui. The hindwing androconial patches 
are much more common and usually lie just outside the cell between 
Ma and Cu 2 , although Mandarinia has a Mycalesis-\ike hair tuft along 
vein Rs (Fig. 73). Hindwing hair tufts are commoner than mealy 
patches, but both are present throughout the tribe. 



LEE D. MILLER 47 

Genera Included in the Lethini 



A eropetes-ser'ies 



Aeropetes Billberg, 1820: 79. Type-species: Papilio tulbaghia Linne, desig- 
nated by Hemming, 1943: 23. 

= Meneris Westwood, 1850, in Doubleday, Westwood and Hewitson, 
1846-1852: 296. Type-species: Papilio tulbaghia Linne, by mono- 
typy. 
Paralethe van Son, 1955: 51. Type-species: Satyrus dendrophilus Trimen, by 
original designation. 

Lethe-series, sensu stricto 

Aphysoneura Karsch, 1894: 190. Type-species: Aphysoneura pigmentaria 
Karsch, by original designation. 

= Rhaphiceropsis Sharpe, 1894: 336. Type-species: Rhaphiceropsis 
pringlei Sharpe (= Aphysoneura pigmentaria Karsch), by original 
designation. 

Archondesa Moore, 1892 (1890-1893): 270. Type-species: Lethe lanaris But- 
ler, by original designation. 

Charma Doherty, 1886: 117. Type-species: Zophoessa baladeva Moore, by 
original designation. Proposed as a subgenus of Lethe Hiibner. 

= Putlia Moore, 1892 (1890-1893): 287. Type-species: Zophoessa 
baladeva Moore, by original designation. Proposed to replace 
Charma Doherty, wrongly believed to be preoccupied. 

Choranesa Moore, 1892 (1890-1893): 270. Type-species: Lethe trimacula 
Leech, by original designation. 

Debis Doubleday and Hewitson, 1849, in Doubleday, Westwood and Hewitson, 
1846-1852: pi. 61. Type-species: Debis samio Doubleday and Hewit- 
son, by monotypy. 

Dionana Moore, 1892 (1890-1893): 271. Type-species: Lethe margaretae 
Elwes, by original designation. 

Enodia Hiibner, [1819] (1816-1826): 61. Type-species: Enodia andromacha 
Hiibner (= Papilio portlandia Fabricius), designated by Scudder, 1872: 
26. 

Hanipha Moore, 1880 (1880-1881): 18. Type-species: Lethe sihala Moore 
(= Lethe dynsate Hewitson), by original designation. This name is 
commonly misspelled "Hanifa". 

Harima Moore, 1892 (1890-1893): 299. Type-species: Neope callipteris But- 
ler, by original designation. 

Hermias Fruhstorfer, 1912 (1912-1915): 324. Type-species: Satyrus verma 
Kollar, by monotypy. Proposed as a "subgenus or species-group" of 
Lethe Hiibner. 

Kerrata Moore, 1892 (1890-1893): 285. Type-species: Lethe tristigmata El- 
wes, by original designation. 

Kirrodesa Moore, 1892 (1890-1893): 237. Type-species: Debis sicelis Hew- 

MEM. AMER. ENT. SOC, 24 



48 



THE SATYRIDAE 




Figs. 60-72. Elymniinae: Lethini. 60. Pararge aegeria (Linne), £ vena- 
tion. 61. P. aegeria, palpus. 62. P. aegeria, $ foreleg. 63. P. aegeria, 2 
foretarsus. 64. Crebeta deidamia (Eversmann), $ venation. 65. Lopinga 
dumetorum (Oberthiir), venation of £ forewing. 66. Aphysoneura pigmentaria 
Karsch, venation of £ forewing. 67. Aeropetes tulbaghia (Linne), cS venation. 
68. A. tulbaghia, palpus. 69. A. tulbaghia, £ foreleg. 70. A. tulbaghia, 2 
foretarsus. 71. A. tulbaghia, midleg. 72. Paralethe dendrophiius (Trimen), 
£ venation. 

itson, by original designation. 
Lethe Hiibner, [1819] (1816-1826): 56. Type-species: Papilio europa Fabri- 
cius, by monotypy. 

= Tanaoptera Billberg, 1820: 79. Type-species: Papilio europa Fab- 



LEE D. MILLER 49 

ricius, designated by Hemming, 1933: 199. 

Magula Fruhstorfer, 1912 (1912-1915): 313. Type-species: Zophoessa jala- 

urida deNiceville, designated by Hemming, 1935: 1. Preoccupied by 

Magula Scudder (Megerle MS.), 1882, but no replacement name has 

been proposed. 

Nemetis Moore, 1892 (1890-1893): 237. Type-species: Papilio minerva Fab- 

ricius, by original designation. 
Neope Butler, 1867d: 166. Type-species: Lasiommata (?) bhadra Moore, 
designated by Butler, 1868b: 112. 

= Enope Moore, 1857: 228. Type-species: Lasiommata (?) bhadra 
Moore, designated by Moore, 1892 (1890-1893): 299. Preoc- 
cupied by Enope Walker, 1854. 
= Blanaida Kirby, 1877: 699. Type-species: Lasiommata (?) bha- 
dra Moore, by original designation. Proposed to replace Enope 
Moore. 
Ninguta Moore, 1892 (1890-1893): 310. Type-species: Pronophila schrenkii 
Menetries, by original designation. 

= A randa Fruhstorfer, 1909: 134. Type-species: Pronophila schrenkii 
Menetries, by original designation. 
Patala Moore, 1892 (1890-1893) : 305. Type-species: Zophoessa yama Moore, 

by original designation. 
Placilla Moore, 1892 ( 1890-1893) : 253. Type-species: Lethe christophi Leech, 

by original designation. 
Ptychandra Felder and Felder, 1861: 304. Type-species: Ptychandra lorquinii 

Felder and Felder, by monotypy. 
Rangbia Moore, 1892 (1890-1893): 232. Type-species: Debis scanda Moore, 

by original designation. 
Samanta Moore. 1880: 166. Type-species: Myca'esis malsara Moore, by 

original designation. 
Satyrodes Scudder, 1875b: 242. Type-species: Papilio eurydice Linne. by 
original designation. 

= Argus Scopoli, 1777: 432. Type-species: Papilio eurydice Linne. 
designated by Scudder, 1872: 27. Preoccupied by Argus Bohadsch. 
1761. 
Sinchula Moore. 1892 (1890-1893): 275. Type-species: Debis sidonis Hew- 

itson, by original designation. 
Tansima Moore, 1881: 305. Type-species: Lethe satyrina Butler, by mono- 
typy. 
Zophoessa Westwood. 1851. in Doubleday, Westwood and Hewitson, 1846- 
1852: 362. Type-species: Zophoessa sura Westwood. by monotypy. 

Pararge-series, 

Chonala Moore, 1893 (1890-1893): 14. Type-species: Debis (Tansima) 
masoni Elwes, by original designation. 

MEM. AMER. ENT. SOC, 24 



50 THE SATYRIDAE 

Crebeta Moore, 1893 (1890-1893): 11. Type-species: Pararge deidamia Evers- 
mann, by original designation. 

Kirinia Moore, 1893 (1890-1893): 14. Type-species: Lasiommata epimen- 
ides Menetries, by original designation. 

Lasiommata Westwood, 1841: 65. Type-species: Papilio megera Linne, desig- 
nated by Scudder. 1875a: 202. 

= Amecera Butler, 1867d: 162. Type-species: Papilio megera Linne, 
designated by Butler, 1868b: 123. 

Lopinga Moore, 1893 (1890-1893): 11. Type-species: Pararge dumetorum 
Oberthiir, by original designation. 

Orinoma Gray, 1846: 14. Type-species: Satyrus (?) damaris Doubleday {nee. 
Gray), by monotypy. 

Pararge Hiibner, [1819] (1816-1826) : 59. Type-species: Papilio aegeria Linne, 
designated by Butler, 1868a: 195. This name has been frequently mis- 
spelled as "Pararga" or "Parage" in the literature. 

Rhaphicera Butler. 1867d: 164. Type-species: Lasiommata satricus Westwood 
and Hewitson, designated by Butler. 1868b: 158. 

Tatinga Moore, 1893 (1890-1893): 5. Type-species: Satyrus thibetanus Ober- 
thiir, by original designation. 

Mandarinia-series 

Mandarinia Leech, 1892: 9. Type-species: Mycalesis regalis Leech, by original 
designation. 

Neorina-series 

Ethope Moore, 1865: 770. Type-species: Mycalesis (?) himachala Moore, by 
monotypy. 

= Theope Moore, 1857: 234. Type-species: Mycalesis (?) hima- 
chala Moore, by original designation. Preoccupied by Theope 
Doubleday and Hewitson, 1847. 
= Anadebis Butler, 1867c: 50. Type-species: Mycalesis (?) hima- 
chala Moore, by monotypy. 
Euploeamima Holland, 1887: 113. Type-species: Zethera diademoides Moore, 

by original designation. 
Hermianax Fruhstorfer, 1912 (1912-1915): 326. Type-species: Neorina lati- 
picta Fruhstorfer (= Neorina lowi Doubleday and Hewitson), by origi- 
nal designation. Proposed as a subgenus of Neorina Westwood. 
Neorina Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852: 
369. Type-species: Neorina hilda Westwood. by original designation. 

Figs. 73-87. Elymniinae: Lethini (Figs. 73-80) and Zetherini (Figs. 81- 
87). 73. Mandarinia regalis (Leech), 6 venation. 74. M. regalis, palpus. 
75. M. regalis, 9 foretarsus. 76. M. regalis, midleg. 77. Neorina hilda West- 
wood, 6 venation. 78. N. hilda, $ foreleg. 79. N. hilda, 2 foretarsus. 80. 



LEE D. MILLER 



51 




N. Hilda, midleg. 81. Zethera pimplea (Erichson), 6 venation. 82. Z. pim- 
ple a, palpus. 83. Z. pimplea, 6 foreleg. 84. Z. pimplea, 2 foretarsus. 85. 
Z. pimplea, midleg. 86. Amechania incerta Hewitson. i venation. 87. Cal- 
large sagitta (Leech). 6 venation. 



MEM. AMER. ENT. SOC. 24 



52 



THE SATYRIDAE 



Tribe Zetherini, new tribe 

(Figures 81-88) 

The zetherines have been considered part of the Elymniini by 
such authors as Gaede (1931), and while these two tribes share such 
features as close forewing veins M- and Cui, they are abundantly 
distinct. Perhaps the closest relatives of the zetherines are certain 
of the more primitive lethines such as Aeropetes, but the differences 
discussed below also serve to separate these two tribes. The pattern 
of the zetherines is highly aberrant: several species are white with 
brown interveinal markings and either no ocelli or poorly developed 
ones, whereas Zethera pimplea (Erichson) looks like the Nearctic 



0.4xML H 

1 WZ/ 




Zethera 



Amechania 



Callarge 



Fig. 88. Elymniinae: Zetherini. Relative lengths of the femur + tibia + 
tarsus of the forelegs (FL, diagonal lines), midlegs (ML. light stippling) and 
hindlegs (HL, heavy stippling) of the genera. In all instances the ML value 
is unity. The top bar for each genus represents the measurements obtained 
from males, the bottom bar those from females. 

Limenitis weidemeyeri (Edwards) (Nymphalidae: Limenitinae), a 
black species with a wide white central band crossing both wings. 

The absence of inflation of the forewing veins serves to distinguish 
this tribe from the Mycalesini, while the spiny midtibia and the well 
developed tibial spurs separate the Zetherini from the Elymniini. 
Most Lethini, with the exception of the Neorina-series, have hairy 
eyes, whereas all zetherines have naked eyes. The relative position 
of the forewing veins M.i, Cui and Cu 2 serve to distinguish members 
of this tribe from the Neorina-series of the Lethini. All the members 
of the Zetherini are Indo- Australian. The zetherines are character- 
ized as follows: 

The eyes are naked. The antennae are just over two-fifths the 



LEE D. MILLER 53 

length of the forewing costa. The antennal club is weakly developed 
and gently tapered, never more than twice the thickness of the shaft. 
The relative lengths of the second and third segments of the palpus 
are variable: in Collar ge the third segment is about one-seventh as 
long as the second, whereas in Zethera it is one-third as long. The 
hairs of the second segment of the palpus are always shorter than the 
segment is wide. 

The male foreleg is well developed with the femur longer than the 
tibia and a mono- to trimerous tarsus which may or may not bear 
spines. The female foreleg is also well developed with a pentamerous, 
slightly clubbed tarsus bearing spines on the first four subsegments. 
The hindlegs are equal to or slightly longer than the midlegs (Fig. 

88). The mid-tibia is long twice to two and a half times as 

long as the first midtarsal subsegment — and the midtibia is spiny 
dorsad, bearing moderately well developed tibial spurs, but no spine 
at the dorsal, distal end. 

The forewing cell is slightly excavate and two-fifths to half as long 
as the forewing costa. The forewing radial veins arise from the cell 
in three branches, and veins Rs and Mi are approximate, but not 
connate, at their origins. Vein M2 arises much nearer Mi than M. f , 
and Cui arises much nearer M3 than Cu 2 . None of the forewing 
veins are much inflated basally. 

The hindwing cell is strongly produced at the origin of M3 and 
is one-third to half the length of the wing measured to the end of 
M3. Hindwing vein Sc + Ri is longer than 3 A. Veins M3 and Cui 
are connate, or nearly so, and M- arises nearer (or in some cases, 
much nearer) Mi than Ma. 

The pattern of all the species is aberrant. Callarge is cream- 
colored with dark veins. Amechania is whitish with dark veins, some 
dark interveinal markings and a few ocelli. Zethera is blackish- 
brown with broad discal white bands on all wings. There are no 
defined androconial patches on any zetherine. 

Genera Included in the Zetherini 

Amechania Hewitson, 1861 (1856-1876): 87. Type-species: Amechania in- 

certa Hewitson, by monotypy. 
Callarge Leech. 1892: 57. Type-species: Zethera sagitta Leech, by original 

designation. 

MEM. AMER. ENT. SOC, 24 



54 



THE SATYRIDAE 



Zethera C. Felder, 1861: 26. Type-species: Cynthia pimplea Erichson, by 
monotypy. 

Tribe Elymniini Herrich-Schaffer, 1864 

(Figures 89-97) 

Eurytelidae Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 

1852: 403 (in part). 
Elymniina Herrich-Schaffer, 1864: 124. 

The Elymniini are a homogeneous group and show some affinities 
within the subfamily. There is little to connect the elymniines with 
the Lethini directly, but the positions of certain of the veins show a 
definite relationship between these butterflies and the zetherines, 
which in turn do relate to the Lethini. The deeply excavate forewing 
cell of these insects is approached in the Mycalesini, and many of the 
latter groups have androconial patches placed very like those which 
characterize the Elymniini. 



0.4xML, FL 




Elymnias 
Elymniopsis 

Melynias 

Mimadelias 

Biudsa 



Fig. 89. Elymniinae: Elymniini. Relative lengths of the femur + tibia 
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and 
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML 
value is unity. The top bar for each genus represents the measurements ob- 
tained from males, the bottom bar those from females. 

This tribe may be distinguished from all others within the sub- 
family by the lack of tibial spurs in all species. Many of the genera 
also have extremely short cells on both wings, but there are a few 
exceptions. 



LEE D. MILLER 55 

All elymniines are Paleotropical, most being found in the Indo- 
Australian region, but a couple of species range as far west as West 
Africa. The Elymniini are characterized in the following paragraphs. 

The eyes are naked. The antennae are about two-fifths as long 
are the forewing costa, although they may be as much as 0.46 times 
as long as the costa. The antennal club is very gentle and seldom 
more than twice the width of the shaft, except in the genus Melynias 
where it is over twice the thickness of the shaft. The third segment 
of the palpus is from one-fifth {Melynias) to one-third (Dyctis) the 
length of the second segment. The hairs of the second palpal seg- 
ment are about as long as the segment is wide. 

The male foreleg is well developed with the femur longer than 
the tibia and with a monomerous, unspined tarsus. The female fore- 
leg is also well developed with a pentamerous, unclubbed tarsus 
bearing a double set of spines on the third and fourth subsegments 
(also a double set on the second subsegment in Elymniopsis) . The 
mid- and hindlegs are of about the same length (Fig. 89). The 
midtibia is generally less than twice the length of the first midtarsal 
subsegment and is smooth dorsad, without tibial spurs or a spine at 
the dorsal, distal end. 

The forewing cell is usually deeply excavate (Figs. 90-92) and 
is always less than half the length of the forewing costa. Frequently 
the cell is less than a third as long as the costa. The forewing radial 
veins arise in three branches from the cell, and Rs and Mi are well 
separated at their origins. Vein M 2 arises much nearer Mi than M.3, 
and Cui is almost connate with M3, never near Cu-. Forewing vein 
Sc is usually inflated basad; the other stems are not. 

The hindwing cell is more or less square-cut because of the dis- 
placement marginad of Mi and is from one-third to almost half as 
long as the wing measured to the end of M3. Hindwing vein Sc + Ri 
is always much shorter than 3A. Veins M3 and Cui are always 
connate, or nearly so, and M2 arises much nearer Mi than M3. 

The pattern is quite variable and has been considered a mimetic 
one. Many of the species resemble various Danaidae, particularly of 
the genus Euploea. For a description of most of the species see 
Moore, 1893 (1890-1893). 

MEM. AMER. ENT. SOC, 24 



56 



THE SATYRIDAE 




Figs. 90-97. Elymniinae: Elymniini. 90. Elymnias hypermnestra (Linne), 
S venation. 91. Bruasa penanga (Westwood and Hewitson), S venation. 
92. Dyctis agondas Boisduval, 6 venation. 93. E. hypermnestra, palpus. 94. 
E. hypermnestra, S foreleg. 95. E. hypermnestra, 9 foretarsus. 96. E. hy- 
permnestra, midleg. 97. Elymniopsis Use Hemming, palpus. 



Genera Included in the Elymniini 

A grusia Moore, 1893 (1890-1893): 144. Type-species: Melanitis esaca West- 
wood and Hewitson, by original designation. 

Bruasa Moore, 1893 (1890-1893): 144. Type-species: Melanitis penanga 
Westwood and Hewitson, by original designation. 

Didonis Hiibner, [1819] (1816-1826): 17. Type-species: Papilio vitellia 
Cramer, designated by Scudder, 1875a: 156. 



LEE D. MILLER 57 

Dyctis Boisduval. 1832: 138. Type-species: Dyctis agondas Boisduval, by 

monotypy. 
Elymnias Hiibner, 1818 (1818-1825): 12. Type-species: Elymnias jynx Hiib- 

ner (= Papilio hypermnestra Linne), designated by Hemming, 1943: 

24. Scudder (1875a: 162) designated Papilio lais Fabricius as type, 

based on Elymnias Hiibner, [1819] (1816-1826): 37. This designation 

was invalidated when the appropriate sections of the "'Zutrage" were 

found to be older than those of the "Verzeichniss". 
Elymniopsis Fruhstorfer. 1907: 171, 173-174. Type-species: Papilio phegea 

Fabricius (= Elymniopsis Use Hemming), designated by Hemming, 

1943: 24. 
Melynias Moore, 1893 (1890-1893): 144. Type-species: Papilio lais Cramer 

(= Papilio nesaea Linne), by original designation. 
Mimadelias Moore, 1893 (1890-1893): 144. Type-species: Elymnias vasu- 

deva Moore, by original designation. 

Tribe Mycalesini, new tribe 
(Figures 98-119) 

The mycalesines show rather close relationships to both the 
Lethini and the Elymniini. Such mycalesine genera as Orsotriaena 
and Bletogona lack the characteristic inflation of the veins, but do 
have the excavate forewing cell and the distinctive androconial 
patches, whereas the lethine Mandarinia regalis (Leech) has the 
androconial patch but lacks the inflation of forewing vein 2A or the 
excavate forewing cell. The excavate forewing cell and the presence 
in some genera of a hair tuft on the upper surface of the hindwing 
at the anterior margin of the cell strongly suggests a relationship 
between the Mycalesini and the Elymniini. 

The characteristic subquadrate shape of the basal inflation of 
forewing vein 2A serves to distinguish the vast majority of all 
mycalesines. In the Hypocystini (Satyrinae) this inflation of 2A 
is suggested, but in that tribe the inflation is in the form of a rounded 
knot, rather than a subquadrate thickening, as in the present tribe. 

The Mycalesini are general throughout the Old World tropics and 
form a homogeneous group. There is some regional correlation of 
genera in the tribe, and for that reason they are arranged geo- 
graphically here. It is interesting to note that although hairy or 
naked eyes do not characterize the tribe, all mainland African Myca- 
lesini have naked eyes, all those from Madagascar have hairy eyes 
and both states are found in the Indo-Australian genera. This 

MEM. AMER. ENT. SOC, 24 



58 



THE SATYRIDAE 



0.4 XML, FL 




Mycalesis 

Satoa 

Orsotriaena 

Bletogona 

Bicyclus 
Dichothyris 

Admiratio 
Heteropsis 

Masoura 



Fig. 98. Elymniinae: Mycalesini. Relative lengths of the femur + tibia 
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and 
hind legs (HL, heavy stippling) of selected genera. In all instances the ML 
value is unity. The top bar for each genus represents the measurements ob- 
tained from males, the bottom bar those from females. 

situation will be discussed further in the section on "Evolution and 
Zoogeography." The Mycalesini are characterized as follows: 

The eyes are naked or hairy: the Indo- Australian species show 
both conditions, those from mainland Africa all have naked eyes 
and those from Madagascar have hairy eyes. The antennae are from 
two-fifths to half the length of the forewing costa. The antennal club 
is moderately well developed, occupying the distal one-third to one- 
fifth of the antenna and thickened from less than twice ( Orsotriaena ) 
to more than three times (such genera as Dalapa and Martanda) the 
thickness of the shaft. The third segment of the palpus is usually 
one-third to one-fifth the length of the second segment, but the third 



LEE D. MILLER 59 

segment is more than half as long as the second in Dalapa. The hairs 
of the second palpal segment are generally from as long as the seg- 
ment is wide to twice the width of the segment. 

The male foreleg is well developed with the femur longer than 
the tibia and one or (rarely) two unspined or weakly spined tarsal 
subsegments. The female foreleg is moderately well developed with 
a pentamerous, lightly clubbed (less so in genera from Madagascar, 
Fig. 118) tarsus bearing spines on at least the fourth subsegment, 
often also on the second and third and occasionally on the first (such 
genera as Martanda) . The mid- and hindlegs are not significantly 
unequal. The midtibia is less than twice as long as the first midtarsal 
subsegment. The midtibia is usually unspined dorsad, although it is 
weakly spined in Orsotriaena (Fig. 108), etc.; tibial spurs are always 
present and well developed and there is no spine at the dorsal distal 
end of the midtibia. 

The forewing cell is excavate and varies little from half the length 
of the forewing costa. The forewing radial veins arise in three 
branches from the cell, and veins Rs and Mi arise separate, but 
approximate, from the cell. Vein M2 arises much nearer Mi than 
Ms, and Cui arises nearer Ma than Cu-. The pattern of basal infla- 
tion of the forewing veins is characteristic, except in Orsotriaena 
(Fig. 106) and Bletogona: veins Sc and the cubital stem are inflated 
normally, but markedly, and 2A is inflated near its base as a sub- 
quadrate knot. 

The hindwing cell is produced at M:s and is one-third to half as 
long as the wing measured to the end of Ms. Hindwing vein Sc + Ri 
is as long as, or shorter than, 3A. Veins Ms and Cui are connate or 
approximate at their origins, except in Culapa (Fig. 104) where they 
are widely separated, and M2 arises much nearer Mi than M3. 

The pattern is highly variable. Many genera show the greatest 
development of the ocelli in forewing space Cui-Cu:>. Most of the 
species are dark brown on both surfaces, hence the common name 
"bushbrowns", but some of the African species are iridescent blue 
or purple above and the Malgache species Masoura masoura (Hewit- 
son) is white with a yellow patch on the forewing, superficially more 
resembling a pierid than a satyrid. The distribution of androconia 
is characteristic: there is a polished area along the inner margin of 
the forewing on the under surface, and other areas of androconial 
concentration are in the area between Cu-2 and 2A of the forewing 

MEM. AMER. ENT. SOC, 24 



60 



THE SATYRIDAE 




Figs. 99-108. Elymniinae: Mycalesini. 99. Mycalesis jrancisca (Stoll), 
$ venation. 100. M. jrancisca, palpus. 101. M. jrancisca, S foreleg. 102. 
M. jrancisca, 2 foretarsus. 103. M. jrancisca, midleg. 104. Culapa mnasicles 
(Hewitson), 8 venation. 105. Mydosama juscum (Felder and Felder), $ 
venation. 106. Orsotriaena medus (Fabricius), 6 venation. 107. O. medus, 
2 foretarsus. 108. O. medus, midleg. 



and along the hindwing cell on both margins, 
frequent than mealy patches in this tribe. 



Hair tufts are more 



Genera Included in the Mycalesini 
Indo-Australian genera 
Bletogona Felder and Felder, 1867 (1864-1867): 465. Type-species: Bleto- 



LEE D. MILLER 61 

gona mycalesis Felder and Felder, by monotypy. 

Calysisme Moore, 1880 (1880-1881): 20. Type-species: Papilio drusia Cra- 
mer, designated by Moore, 1891 (1890-1893): 172. 

Celebina Fruhstorfer, 1898: 263. Type-species: Celebina inga Fruhstorfer, 
designated by Hemming, 1935: 1. 

Culapa Moore, 1878: 825. Type-species: Mycalesis mnasicles Hewitson, by 
monotypy. 

Dolapa Moore, 1880: 158. Type-species: Mycalesis sudra Felder, by original 
designation. 

Hamadryopsis Oberthiir, 1899: 17. Type-species: Hamadryopsis drusillodes 
Oberthiir, by monotypy. 

= Drusillopsis Fruhstorfer, 1908: 217. Type-species: Hamadryopsis 
drusillodes Oberthiir, by monotypy. 

Indalasa Moore, 1880: 166. Type-species: Mycalesis moorei Felder, by mono- 
typy. 

Jatana Moore, 1880: 164. Type-species: Mycalesis mynois Hewitson, by 
monotypy. 

Kabanda Moore, 1880: 168. Type-species: Mycalesis malsarida Butler, by 
original designation. 

Loesa Moore, 1880: 177. Type-species: Mycalesis oroatis Hewitson, by mono- 
typy. 

Lohora Moore. 1880: 175. Type-species: Mycalesis dexamenus Hewitson, by 
original designation. 

Martanda Moore, 1880: 169. Type-species: Mycalesis janardana Moore, by 
original designation. 

Monotrichtus Hampson, 1891: 179. Type-species: Mycalesis safitza Hewitson, 
by original designation. Originally proposed as a subgenus of Myca- 
lesis Hiibner. 

Mycalesis Hiibner, 1818 (1818-1825): 17. Type-species: Papilio francisca 
Stoll, designated by Hemming, 1937: 149. Butler, 1868a: 196, desig- 
nated Papilio evadne Cramer as type, based on Mycalesis Hiibner, 
[1819] (1816-1826): 55. This designation was invalidated when the 
appropriate sections of the "Zutrage" were found to be older than those 
of the "Verzeichniss". 

= Gareris Moore, 1880: 156. Type-species: Mycalesis sanatana 
Moore (= Papilio francisca Stoll), by original designation. 

Mydosama Moore, 1880: 170. Type-species: Dasyomma fuscum Felder and 
Felder, by original designation. 

= Dasyomma Felder and Felder, 1860: 401. Type-species: Dasy- 
omma fuscum Felder and Felder, by monotypy. Preoccupied by 
Dasyomma Macquart, 1841. 

Myrtilus deNiceville, 1891: 341. Type-species: Mycalesis (Myrtilus) mystes 
deNiceville, by original designation. Proposed as a subgenus of Myca- 
lesis Hiibner. 



MEM. AMER. ENT. SOC, 24 



62 



THE SATYRIDAE 




Figs. 109-119. Elymniinae: Mycalesini. 109. Bicyclus hewitsonii (Dou- 
met), $ venation. 110. B. hewitsonii, palpus. 111. B. hewitsonii, $ foreleg. 
112. B. hewitsonii, ? foretarsus. 113. Dichothyris sambulos (Hewitson), $ 
venation. 114. Heteropsis drepana Westwood, antennal club. 115. H. dre- 
pana, $ venation. 116. H. drepana, palpus. 117. H. drepana, S foreleg. 
118. H. drepana, 2 foretarsus. 119. Admiratio paradoxa (Mabille), $ vena- 
tion. 

Nasapa Moore, 1880: 176. Type-species: Mycalesis aramis Hewitson, by 
monotypy. 

Nebdara Moore, 1880: 173. Type-species: Mycalesis tagala Felder, by origi- 
nal designation. 

Nissanga Moore, 1880: 169. Type-species: Mycalesis patnia Moore, by origi- 
nal designation. 

Orsotriaena Wallengren, 1858: 79. Type-species: Papilio medus Fabricius ( = 
Papilio hesione Cramer), by monotypy. 

Pachama Moore, 1880: 165. Type-species: Mycalesis mestra Hewitson, by 
monotypy. 



LEE D. MILLER 63 

Physcon deNiceville, 1898: 133. Type-species: Mycalesis pandoea Hopffer, by 
original designation. Proposed as a subgenus of Mycalesis Hubner. 

Sadarga Moore, 1880: 157. Type-species: Mycalesis gotama Moore, by origi- 
nal designation. 

Samundra Moore, [1890-1892] (1890-1893): 162. Type-species: Mycalesis 
anaxioides Marshall and deNiceville, by monotypy. 

Satoa Moore, 1880: 157. Type-species: Mycalesis maianeas Hewitson, by 
monotypy. 

Sevanda Moore, 1880: 174. Type-species: Satyrus duponcheli Guerin, by 
original designation. 

Suralaya Moore, 1880: 159. Type-species: Mycalesis oreseis Hewitson, by 
monotypy. 

Telinga Moore, 1880: 167. Type-species: Satyrus adolphei Guerin, by mono- 
typy. 

Virapa Moore, 1880: 156. Type-species: Mycalesis anoxias Hewitson, by origi- 
nal designation. 

African genera 

Bicyclus Kirby, 1871a: 47. Type-species: Idiomorphus hewitsonii Doumet, 
designated by Kirby, 1871b: 363. 

= Idiomorphus Doumet, 1861: 174. Type-species: Idiomorphus 
hewitsonii Doumet, by monotypy. Preoccupied by Idiomorphus 
Chaudoir, 1846. 
Dichothyris Karsch, 1893: 203. Type-species: Mycalesis sambulos Hewitson, 

designated by Hemming, 1935: 1. 
Hallelesis Condamin, 1960: 1257. Type-species: Mycalesis asochis Hewitson. 
designated by Condamin, 1961: 783. 

Malgache genera 

Admiratio Hemming, 1964b: 137. Type-species: Smithia paradoxa Mabille, 
by original designation. 

= Smithia Mabille, 1880: 173. Type-species: Smithia paradoxa 
Mabille, by monotypy. Preoccupied by Smithia Milne Edwards 
and Haime, 1851. 
Henotesia Butler, 1879: 228. Type-species: Henotesia wardii Butler, by origi- 
nal designation. 
Heteropsis Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 
1852: 323. Type-species: Heteropsis drepana Westwood, by monotypy. 
Houlbertia Oberthur, 1916: 199. Type-species: Erebia passandava Ward, des- 
ignated by Hemming, 1964a: 120. 
Masoura Hemming, 1964b: 138. Type-species: Melanitis masoura Hewitson, 
by original designation. 

= Gallienia Oberthur, 1916: 205. Type-species: Melanitis masoura 
Hewitson, designated by Hemming, 1964a: 120. 

MEM. AMER. ENT. SOC, 24 



64 



THE SATYRIDAE 



Subfamily Eritinae, new subfamily 

It is obvious that the two genera in this subfamily are intermediate 
between the Elymniinae (and closest to the Lethini of that subfamily) 
and the Satyrinae, but perhaps the present genera should not be com- 
bined into a single sub-family or tribe. Nevertheless, these genera 
seem closest to one another, and I should be reluctant to erect two 
monotypic taxa for them. 

The female foretarsus is pentamerous and unclubbed (Figs. 125, 
131) and more primitive than that of the Satyrinae. The pattern 
is rather more primitive than that of most satyrines (Schwanitsch, 
1924) and more closely approximates the pattern of the lethines. 
The male foreleg of the genus Coelites is much reduced over the 
situation in the lethines, however, and the forewing cell of Erites 
is much longer than that of any elymniine. 

Both genera are found on the Malay peninsula and in Indonesia. 
A single tribe, the Eritini, includes all species. 

Tribe Eritini, new tribe 

(Figures 120-132) 

The diagnostic features of the subfamily Eritinae and of the only 
tribe, the Eritini, are as follows: 

The eyes are naked. The antennae are about two-fifths the length 



0.4* ML 




Erites 



Coelites 



Fig. 120. Eritinae: Eritini. Relative lengths of the femur + tibia + tar- 
sus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and 
hindlegs (HL, heavy stippling) of the genera. In all instances the ML value 
is unity. The top bar for each genus represents the measurements obtained 
from males, the bottom bar those from females. 

of the forewing costa. The antennal club is indistinct {Coelites, 
Fig. 122) to gradual (Erites, Fig. 128) and never more than twice 
as thick as the shaft. The third segment of the palpus is about 



LEE D. MILLER 65 

one-fifth as long as the second, and the development of the hairs of 
the second segment is variable: in Coelites they are about as long as 
the segment is wide, whereas in Erites the hairs are two and a half 
times the width of the segment. 

The male foreleg is variably developed: that of Erites is slightly 
miniaturized with subequal femur and tibia and a monomerous, 
unspined tarsus (Fig. 130); whereas that of Coelites is greatly 
reduced, the femur longer than the tibia and the tarsus represented 
only by a bump at the tip of the tibia (Fig. 124). The female foreleg 
is moderately well developed with a pentamerous, unclubbed tarsus 
bearing spines on the third and fourth subsegments {Erites, Fig. 131) 
or on the first four subsegments (Coelites, Fig. 125). The midlegs 
are approximately equal to, or slightly longer than, the hindlegs 
(Fig. 120). The midtibia is just over twice as long as the first 
midtarsal subsegment and is smooth dorsad with moderately well 
developed tibial spurs and no spines at the dorsal, distal end. 

The forewing cell is produced at the origin of Mi and one-half 
{Coelites, Fig. 121) to three-fifths (Erites, Fig. 127) as long as the 
forewing costa. The forewing radial veins arise in three branches 
from the cell, and Rs and Mi are approximate. Vein M-- arises 
slightly nearer (Erites) to much nearer (Coelites) Mi than M 3 , and 
Cui arises somewhat nearer M.3 than Cuo. Only forewing vein Sc 
is inflated basad. 

The hindwing cell is produced at the origin of Ms and is less 
than half (Coelites, Fig. 121) to three-fifths (Erites, Fig. 127) as 
long as the wing measured to the end of Ms. Hindwing vein Sc + Ri 
is longer than 3 A, and veins M 3 and Cui may be connate (Coelites) 
to widely separated (Erites) at their origins. Vein Mi; arises slightly 
nearer Mi than Ms. 

Erites is an olive-brown insect above with the lines of the under 
surface faintly indicated and ocelli in forewing spaces M1-M2, M2-M3 
and a large one in Cui-Cu^ and hindwing spaces Rs-Mi through 
CU1-CU2. On the under surface the pattern is the same but there 
are dark striatums and transverse ochre lines in the medial areas of 
both wings. Coelites is uniform gray-brown above with a basal 
purplish sheen and a patch of oily hairs between hindwing veins 2A 
and 3A. The under surface is dark brown basad, gray-brown mar- 
MEM. AMER. ENT. SOC, 24 



66 



THE SATYRIDAE 




Figs. 121-132. Eritinae: Eritini. 121. Coelites nothis Westwood and 
Hewitson, $ venation. 122. C. nothis, antennal club. 123. C. nothis, palpus. 
124. C. nothis, S foreleg. 125. C. nothis, 9 foretarsus. 126. C. nothis, mid- 
leg. 127. Erites madura (Horsefield), S venation. 128. E. madura, antennal 
club. 129. E. madura, palpus. 130. E. madura, S foreleg. 131. E. madura, 
$ foretarsus. 132. E. madura, midleg. 

ginad with ocelli in hindwing spaces Rs-Mi through Cui-Cul.. 



Genera Included in the Eritini 

Coelites Westwood and Hewitson, 1850, in Doubleday, Westwood and Hewit- 
son, 1846-1852: pi. 66. Type-species: Coelites nothis Westwood and 
Hewitson, designated by Butler, 1868a: 195. 

Erites Westwood, 1850, in Doubleday, Westwood and Hewitson, 1846-1852: 
392. Type-species: Hipparchia madura Horsefield, by monotypy. 



LEE D. MILLER 



67 



Subfamily Ragadiinae Herrich-Schaffer, 1864 
Ragadiina Herrich-Schaffer, 1864: 124. 

It is difficult to place the Ragadiinae in a satisfactory systematic 
position, but for reasons to be enumerated below, this subfamily 
appears to occupy a position intermediate between the Elymniinae 
and the Satyrinae. In common with the former, hindwing veins 
M:s and Cui arise from a common point on the cell, but the abortion 
of the forelegs of both sexes (Figs. 136, 137) is a characteristic 
shared with the Satyrinae and Coelites of the Eritinae. In other 
respects, however, the ragadiines stand apart from all of the other 
subfamilies. 

The configuration of the hindwing cell (Figs. 134, 139) is dis- 
tinctive. In males the cell is closed by narrow vestigial veins, whereas 
in the females of some species the cell is completely open between the 
origins of veins M2 and M 3 . 

All members of the Ragadiinae are native to the Indo-Australian 
region and constitute a single tribe, the Ragadiini. Rober (1892) 
split this subfamily, referring Acrophtalmia to the Ypthima group 
and Ragadia to the Mycalesis group, but the characteristic venation 
of the hindwing cannot be ascribed merely to the convergence of two 
different phyletic lines. 

Tribe Ragadiini Herrich-Schaffer, 1864 
(Figures 133-139) 
Ragadiina Herrich-Schaffer, 1864: 124. 



0.2xML 




Ragadia 
Acrophtalmia 



Fig. 133. Ragadiinae: Ragadiini. Relative lengths of the femur + tibia 
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and 
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML 
value is unity. The top bar for each genus represents the measurements ob- 
tained from males, the bottom bar those from females. 

MEM. AMER. ENT. SOC, 24 



68 



THE SATYRIDAE 




Figs. 134-139. Ragadiinae: Ragadiini. 134. Ragadia crisia (Geyer), $ 
venation. 135. R. crisia, palpus. 136. R. crisia, 8 foreleg, showing also the 
trochanter and the distal portion of the coxa. 137. R. crisia, 9 foretarsus. 
138. R. crisia, midleg. 139. Acrophtalmia artemis Felder and Felder, S vena- 
tion. 

The diagnostic features of the subfamily Ragadiinae and its only 
tribe, the Ragadiini, are as follows: 

The eyes are naked. The antennae are no more than half the 
length of the forewing costa. The antennal club is moderately well 
developed, occupies the distal one-fourth to two-fifths of the antenna 
and is thickened to two and a half to three times the width of the 
shaft. The third segment of the palpus is from less than a quarter 
(Ragadia) to slightly less than one-third (Acrophtalmia) the length 
of the second segment. The hairs of the second segment of the 
palpus are shorter than the segment is wide. 

The male foreleg is greatly reduced with but a bump representing 
the tarsus in some species (Ragadia, Fig. 136, for example); the 
tarsus is completely fused with the tibia and indistinguishable from 
it in Acrophtalmia. The femur and tibia are of about the same 
length. The female foreleg is also miniaturized and the tarsus clubbed 
with five subsegments bearing spines on the first four. The mid- and 
hindlegs are of about the same length. The midtibia is less than 
twice the length of the first midtarsal subsegment, is smooth dorsad 
and bears weakly developed tibial spurs, but no spine at the dorsal, 
distal end. The relative lengths of the legs are shown in Fig. 133. 

The forewing cell is excavate near the origin of Ml> and about 



LEE D. MILLER 69 

three-fifths the length of the forewing costa. The forewing radial 
veins arise in two branches from the cell, and veins Rs and Mi are 
nearly connate. Vein M2 arises nearer Mi than M3, and Cm arises 
much nearer M.h than Cu:>. Forewing vein Sc is inflated basad, but 
the other veins are not. 

The hindwing cell is distinctive. It is open in some females 
and at most closed by small, aberrant veins (Figs. 134, 139). The 
cell is from less than half {Acrophtalmia) to more than three-fifths 
(Ragadia) as long as the wing measured to the end of M3. The 
relative lengths of hindwing veins Sc + Ri and 3 A vary: Sc + Ri is 
longer in Acrophtalmia, and 3 A is longer in Ragadia. Veins Mm and 
Cui are connate, whereas M2 arises basad of Mi and much closer to 
it than to M.i. 

The pattern is highly modified, either brown with a broad central 
white band or with alternating brown and off-white stripes. The 
ocelli are poorly developed, except in Ragadia which has a full com- 
plement on the under surface. An androconial hair tuft lies on the 
aberrant veins of the hindwing cell (Figs. 134, 139) and is charac- 
teristic of the tribe. 

Genera Included in the Ragadiini 

Acrophtalmia Felder and Felder, 1861: 305. Type-species: Acrophtalmia ar- 
temis Felder and Felder, by monotypy. Many authors, following Fel- 
der and Felder, 1867 (1864-1867): 486, have misspelled this name 
" Anophthalmia" '. 
Acropolis Hemming, 1934b: 77. Type-species: Acrophthalmia (sic.) thalia 
Leech, by original designation. 

= Pharia Fruhstorfer, 1912 (1912-1915): 295. Type-species: Ac- 
rophthalmia (sic.) thalia Leech, by monotypy. Preoccupied by 
Pharia Gray, 1840. 
Ragadia Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852: 
376. Type-species: Euptychia crisia Geyer, by monotypy. Proposed as 
a subgenus of Neonympha Hiibner. 

Subfamily Satyrinae Boisduval, 1836 

Satyrides Boisduval, 1836: 166; Satyridae Swainson, 1840: 86. 
= Hipparchiadae Kirby, 1837: 297. 
= Maniolinae Hampson, 1918: 385. 

The Satyrinae are the most evolved members of the family and 

MEM. AMER. ENT. SOC, 24 



70 



THE SATYRIDAE 



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MEM. AMER. ENT. SOC, 24 



72 THE SATYRIDAE 

have become the dominant group throughout most of the world. 
Fairly close relationships with three other subfamilies, the Elymniinae, 
Eritinae and Ragadiinae, are indicated by analysis of structures, and 
there are three instances of gradation between the elymniines and the 
satyrines. There is probably only convergence between the Aero- 
petes-series of the Lethini and the Dirini of the present subfamily. 
Far more significant evolutionarily, however, are such genera as 
Lamprolenis in the primitive satyrine tribe Hypocystini. These but- 
terflies show certain similarities to some Mycalesini and Lethini 
(Mandarinia) of the Elymniinae, particularly as regards pattern and 
the placement of the androconial areas. This is probably the stock 
from which the Satyrinae evolved. The Lethini are connected to 
the Satyrinae through the sub-family Eritinae, as indicated in the 
discussion of the eritines. This evidence seems to place the Lethini 
as the closest present-day ancestral stock of the Satyrinae. The 
relationship with the ragadiines is obscure and more in the nature of 
an association because of the greatly reduced forelegs of both sexes. 

Any satyrid with fewer than five subsegments on the female fore- 
tarsus is a member of this subfamily. Even when there are five 
subsegments on the female foreleg of a member of the present sub- 
family, the tarsus is formed into a clublike structure totally unlike 
anything found in the Haeterinae, Brassolinae, Biinae, Elymniinae or 
Eritinae, but hinted at in the Ragadiinae. Comparative female fore- 
tarsi of the pentamerous type are shown in Figs. 5 (Haeterinae), 15 
(Brassolinae), 24, 32 (Biinae), 52, 84, 95, 102 (Elymniinae), 131 
(Eritinae), 137 (Ragadiinae) and 145, 174, 200 (Satyrinae). A 
greatly reduced male foreleg will serve to separate the Satyrinae from 
all other satyrid subfamilies except the Ragadiinae and Coelites of the 
Eritinae. None of the satyrines have the hindwing veins Ms and Cui 
connate, nor the open hindwing cell of the ragadiines. The hindwing 
cell, measured to the origin of vein M 3 , is greater than half as long 
as the wing to the end of Ms, except in some Dirini. The antennal 
club is always moderately well and frequently highly developed. 

The Satyrinae are virtually cosmopolitan, with representatives 
in all parts of the world where butterflies are found, except some 
oceanic islands. Unlike the other subfamilies, the Satyrinae are 
highly developed in temperate regions. 

Ten tribes are recognized in the Satyrinae. The most primitive 



LEE D. MILLER 73 

tribe, the Hypocystini, contains genera which span the gap between 
the other two groups. The Ypthimini-section, characterized more or 
less by their smaller size, frailer appearance and an ecological prefer- 
ence for grasslands, contains the tribes Ypthimini, Euptychiini, Coe- 
nonymphini, Maniolini and Erebiini. The Satyrini-section, larger 
and sturdier insects with woodland or scrubland preferences, is com- 
posed of the tribes Dirini, Pronophilini, Satyrini and Melanargiini. 
A key to the tribes of the Satyrinae is given below, and a diagnostic 
chart of these tribes is provided in Table 6. 

Key to the Tribes of the Satyrinae 

1 . Hindwing cell half or less the length of the wing; forewing cell rounded 

distad (Figs. 248, 255, 256); South Africa Dirini 

Hindwing cell greater than half the length of the wing; forewing cell 
square-cut or excavate 2 

2. Midtibia with a long spine at the dorsal distal end (Fig. 301); Holarc- 

tic most Satyrini 

Midtibia without such a spine 3 

3. Hindwing cell prolonged by a distad displacement of crossvein m=-m.-i 

along vein M= (Fig. 261, for example) 4 

Hindwing cell more or less square cut with crossvein rrb-m.! in line with 
mi-rm (Fig. 309, for example) 8 

4. Female foretarsus with fewer than five subsegments 5 

Female foretarsus pentamerous 6 

5. Third segment of palpus less than one-fourth the length of the second 

(Fig. 305); Holarctic Satyrini (Oenm-series) 

Third segment of palpus longer than one-fourth the length of the second 
(Fig. 282); Neotropical some Pronophilini 

6. Midtibia spiny; Neotropics most Pronophilini 

Midtibia smooth 7 

7. Neotropical species a few Pronophilini 

Australian region species Hypocystini 

8. White (or cream-colored) species, marbled with 

black: Palearctic Melanargiini 

Generally brown or orange species, if white 

not marbled 9 

9. Female foretarsus with fewer than five subsegments 10 

Female foretarsus pentamerous 12 

10. Forewing veins only slightly inflated (Figs. 

234, 240); Holarctic Erebiini 

Forewing veins inflated (Fig. 224, for example) 11 

MEM. AMER. ENT. SOC, 24 



74 THE SATYRIDAE 

11. Midtibia twice or more the length of the first 

midtarsal subsegment; Holarctic Coenonymphini 

Midtibia less than twice as long as the first 

midtarsal subsegment; Holarctic Maniolini 

12. Forewing with a double-pupilled ocellus in space 

Mi-M a or Mi-Ma; Old World 13 

Forewing with no ocellus in space Mi-M.i, or if 

present only single-pupilled; New World Euptychiini 

13. Eyes naked Ypthimini 

Eyes hairy genus Palaeonympha 

Tribe Hypocystini, new tribe 
(Figures 140-169) 

The Hypocystini are here considered to be the most primitive 
members of the Satyrinae, primarily because of the configurations 
of the male and female forelegs. Although these butterflies are not 
closely allied to any other tribe within the Satyrinae, they seem to 
connect the Ypthimini- and Satyrini-sections. The Xenica-series in 
the present tribe connects with the Satyrini-section by pattern and 
structural similarities, whereas the Hypocysta-series exhibits an even 
closer approach to the Ypthimini-section. The New Guinean hypo- 
cystine genus Lamprolenis shows many characteristics in common 
with the elymniine tribes Mycalesini and Lethini. This genus has the 
typical mycalesine androconial patch along the anterior portion of 
the hindwing, and the general facies are reminiscent of the mycalesines 
or of Mandarinia. This similarity may offer an explanation of the 
evolution of the Satyrinae and will be discussed in greater detail later. 

Parallel evolution is shown very well by the New Zealand mem- 
bers of this tribe, with the exception of Dodonidia. The other 
members, living in montane situations, have evolved into butterflies 
very like the Holarctic genus Erebia — indeed, several were placed in 
that genus by early authors. Because the New Zealand "coppers" are 
placed in the Holarctic genus Lycaena Fabricius (H. K. Clench, per- 
sonal communication), the possibility that Argyrophenga and espe- 
cially Erebiola and Percnodaimon are indeed referable to Erebia 
had to be examined, but there is no morphologic evidence to support 
their inclusion in the Holarctic genus. The New Zealand "Erebia" 
are highly modified Hypocystini, most nearly related to the Australian 
members of the tribe. 



LEE D. MILLER 



75 




Hypocysta 

Platypthima 
Zipaetis 
Hyalodia 

Xenica 

Heteronympha 

Dodonidia 

Argynnina 

Argyrophenga 



Fig. 140. Satyrinae: Hypocystini. Relative lengths of the femur + 
tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip- 
pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances 
the ML value is unity. The top bar for each genus represents the measurements 
obtained from males, the bottom bar those from females. 

Members of the New Guinean hypocystine fauna (Platypthima, 
Erycinidia, Piehdopsis and Lamprolenis) have evolved into bewil- 
dering butterflies. In fact, in wing shape and general appearance 
Erycinidia is a classic riodinid, and Pieridopsis is a fine pierid 
"mimic". 

The less miniaturized forelegs in both sexes serves to distinguish 
the hypocystines from all other Satyrinae. The forewing cell is usually 
deeply excavate in this tribe, a characteristic shared with such 
Ypthimini-section tribes as the Ypthimini and Coenonymphini, but 
the hindwing crossvein mrraa is produced marginad along M 2 , as 
in much of the Satyrini-section. Forewing vein 2A is usually inflated 
basad, recalling the Mycalesini, but the inflated part is not sub- 
MEM. AMER. ENT. SOC, 24 



76 



THE SATYRIDAE 



quadrate in the hypocystines. A few of the Xenica-series have little 
or no inflation of forewing vein 2A. 

All members of the Hypocystini are found in the Australian 
region, a few being found as far west as Indonesia. Most of the 
species are found in New Zealand, Australia, New Guinea and the 
associated oceanic islands. Two series of genera are recognized. 




Figs. 141-149. Satyrinae: Hypocystini. 141. Hypocysta euphemia West- 
wood and Hewitson, $ venation. 142. H. euphemia, antennal club. 143. H. 
euphemia, palpus. 144. H. euphemia, S foreleg. 145. H. euphemia, 2 fore- 
tarsus. 146. H. euphemia, midleg. 147. Erycinidia gracilis Rothschild and 
Jordan, 6 venation. 148. Lamprolenis nitida Godman and Salvin, S vena- 
tion. 149. Pieridopsis virgo Rothschild and Jordan, 6 venation. 

The Hypocy sta-sches, sensu stricto, most of which are exclusively 
tropical insects, are characterized by the generally absent tibial spurs, 



LEE D. MILLER 



77 



the longer third segment of the palpus and hindwing vein Ma arising 
nearer M3 than Mi. 

The Xenica-series is restricted to the temperate part of the region 
and characterized by the development of the tibial spurs, the shorter 
third segment of the palpus and (in many genera) hindwing vein 
M-, which arises midway between Mi and M3. 

The Hypocystini are characterized as follows: 

The eyes are generally naked, although they are hairy in such 
diverse genera as Argynnina, Heteronympha, Nesoxenica and Pla- 
typthima. The antennae are generally two-fifths to half the length 
of the forewing costa, but the antennae of Argyronympha are three- 
fifths the length of the costa. The antennal club is well developed in 
most genera, occupying the distal one-fifth to one-third of the antenna 
and expanded to three to four times the width of the shaft (just over 
twice the width of the shaft in Lamprolenis) . The club is eccen- 
trically developed in some New Zealand genera (Figs. 163, 166, 
1 69 ) . The third segment of the palpus is variously one-half to 
one-third the length of the second in the Hypocysta-series (Fig. 143) 
and one-fourth to one-fifth the length of the second in the Xenica- 
series (Fig. 157). The hairs of the second segment of the palpus 
vary from less than the width of the segment to more than twice as 
long as the segment is wide in the Hypocysta-series to over four times 
the width of the segment in some of the New Zealand genera. 





Figs. 150-155. Satyrinae: Hypocystini, all Zipaetis saitis Hewitson. 150. 
$ venation. 151. antennal club. 152. palpus. 153. $ foreleg. 154. 2 
foretarsus. 155. midleg. 



MEM. AMER. ENT. SOC, 24 



78 THE SATYRIDAE 

The male foreleg is moderately well developed (Fig. 140); the 
tibia is usually longer than the femur, and the tarsus is usually mono- 
merous, but may have as many as four subsegments. The female 
foreleg is also well developed (Fig. 140) with a pentamerous, only 
slightly clubbed tarsus bearing spines on the first four subsegments 
(Fig. 145, for example) — there are more than one pair of spines 
on each of the first four subsegments in Dodonidia (Fig. 164). The 
midlegs are as long as, or longer than, the hindlegs (Fig. 140). The 
midtibia is from less than twice to more than twice the length of the 
first midtarsal subsegment, and the tibia is smooth dorsad. Tibial 
spurs are generally well developed in the Xenica-series (Fig. 160), 
including the New Zealand genera; but the spurs are generally absent 
in the Hypocy st a-series (Fig. 146), but they are weakly developed 
in Pieridopsis and moderately developed in Zipaetis (Fig. 155). No 
spine is developed on the dorsal, distal end of the midtibia. 

The forewing cell is excavate, often deeply so, and is from more 
than half to more than three-fifths the length of the forewing costa. 
In the Hypocysta-series, Argyrophenga and Dodonidia the forewing 
radial veins arise in two branches from the cell; in the other genera 
the radials arise in three branches. Forewing veins Rs and Mi are 
connate in Argyrophenga, Erebiola and Percnodaimon, and separate, 
though often approximate, in the other genera. Vein 1VL> arises 
nearer, often much nearer, Mi than Ms, and Cui arises a little nearer 
M:s than Cun. Vein Sc is characteristically inflated in all genera but 
Dodonidia (Fig. 162), and the cubital stem and 2A are inflated in 
such diverse genera as Argyrophenga (Fig. 167), Xenica (Fig. 156), 
Geitoneura (Fig. 161) and most of the Hypocysta-series (Fig. 141, 
for example ) . 

The hindwing cell is usually produced at m 2 -m3, but it is more 
blunted in Dodonidia (Fig. 162) and Zipaetis (Fig. 150), and the 
cell is usually half to three-fifths the length of the wing measured to 
the origin and end of M 3 , respectively. The relative lengths of the 
hindwing veins Sc + Ri and 3 A are variable. Veins M3 and Cui arise 
separately, and M 2 commonly arises midway between Mi and Ms, 
but nearer the latter in the New Zealand "Erebia" and the Hypocysta- 

Figs. 156-169. Satyrinae: Hypocystini. 156. Xenica achanta (Donovan), 
$ venation. 157. X. achanta, palpus. 158. X. achanta, £ foreleg. 159. X. 
achanta, 2 foretarsus. 160. X. achanta, midleg. 161. Geitoneura klngii 



LEE D. MILLER 



79 




(Guerin), 8 venation. 162. Dodonidia helmsii Butler, S venation. 163. D. 
helmsii, antennal club. 164. D. helmsii, 9 foretarsus. 165. Percnodaimon 
pluto (Fereday), $ venation. 166. P. pluto, antennal club. 167. Argy- 
rophenga antipodum Doubleday, $ venation. 168. Erebiola buried Fereday, 
$ venation. 169. E. butleri, antennal club. 



MEM. AMER. ENT. SOC, 24 



80 THE SATYRIDAE 

series. 

The pattern is variable and highly modified. Many of the species 
are illustrated by Waterhouse and Lyell (1914) and by Fruhstorfer 
(191 2[ 191 2- 191 5]). There is a remarkable resemblance between 
Percnodaimon, Erebiola and Argyrophenga and the Holarctic Erebia. 

Genera Included in the Hypocystini 
Hypocysta-series, sensu stricto 

Argyronympha Mathew, 1886: 346. Type-species: Argyronympha pulchra 
Mathew, designated by Hemming, 1943: 23. 

Erycinidia Rothschild and Jordan, 1905: 457. Type-species: Erycinidia gra- 
cilis Rothschild and Jordan, by monotypy. 

Harsiesis Fruhstorfer, 1912(1912-1915): 299. Type-species: Hypocysta hy- 
geia Hewitson, by monotypy. 

Hyalodia Jordan, 1924: 285. Type-species: Hypocysta tenuisquamosa Joicey 
and Talbot, by monotypy. 

Hypocysta Westwood and Hewitson, 1850, in Doubleday, Westwood and 
Hewitson, 1846-1852: pi. 67. Type-species: Hypocysta euphemia 
Westwood and Hewitson, by monotypy. 

Lamprolenis Godman and Salvin, 1880: 610. Type-species: Lamprolenis 
nitida Godman and Salvin, by monotypy. 

Pieridopsis Rothschild and Jordan, 1905: 457. Type-species: Pieridopsis virgo 
Rothschild and Jordan, by original designation. 

Platypthima Rothschild and Jordan, 1905: 458. Type-species: Platypthima 
ornata Rothschild and Jordan, by original designation. 

Zipaetis Hewitson, 1863(1856-1876): [100]. Type-species: Zipaetis saitis 
Hewitson, designated by Butler, 1868a: 194. This genus is exception- 
ally aberrant and is only provisionally placed here. 

Xenica-series 

Argynnina Butler, 1867d: 165. Type-species: Lasiommata hobartia Westwood 
and Hewitson, designated by Butler, 1868a: 196. 

Argyrophenga Doubleday, 1845: 307. Type-species: Argyrophenga antipodum 
Doubleday, by monotypy. 

Dodonidia Butler, 1884: 172. Type-species: Dodonidia hehnsii Butler, by 
original designation. 

Erebiola Fereday, 1879: 128. Type-species: Erebiola butleri Fereday, by 
original designation. 

Geitoneura Butler, 1867d: 164. Type-species: Satyrus klugii Guerin, desig- 
nated by Butler, 1868b: 166. 



LEE D. MILLER 81 

Heteronympha Wallengren, 1858: 78. Type-species: Papilio merope Fabricius, 
designated by Butler, 1868a: 195. 

=Hipparchioides Butler, 1867c: 125. Type-species: Papilio merope 
Fabricius, designated by Butler. 1868b: 99. 
Nesoxenica Waterhouse and Lyell, 1914: 35. Type-species: Lasiommata 
leprea Hewitson, by original designation. 

= Xeniconympha Noricky, 1923: 60. Type-species: Lasiommata 
leprea Hewitson, by original designation. 
Oreixenica Waterhouse and Lyell, 1914: 41. Type-species: Lasiommata (?) 

lathionella Westwood, by original designation. 
Paratisiphone Watkins, 1928: 615. Type-species: Lasiommata lyrnessa Hewit- 
son, by original designation. 
Percnodaimon Butler, 1876: 152. Type-species: Erebia pluto Fereday, by 

original designation. 
Tisiphone Hiibner, [ 1 8 1 9] ( 1816-1826) : 60. Type species: Papilio abeona 

Donovan, designated by Butler, 1868b: 71. 
Xenica Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852: 
387. Type-species: Papilio achanta Donovan, designated by Scudder. 
1875a: 289. 

Tribe Ypthimini, new tribe 
(Figures 170-188) 

This tribe lies at the base of the Ypthimini-section of the Satyrinae 
and seems in some respects to connect with the Hypocysta-sevies of 
the last tribe. In its turn the Ypthimini appears to have given rise 
to the Holarctic Erebiini and Coenonymphini and the Neotropical 
Euptychiini; the Maniolini were evolved later from a coenonymphine 
stock. 

The Ypthimini are generally characterized by a pattern feature: 
the forewing ocellus in space M1-M3 has a double pupil of white or 
bluish-white scales. Forewing vein 2A is not inflated in the ypthi- 
mines, whereas in the Coenonymphini and many Euptychiini this 
vein is greatly distended. While the hindwing crossvein ms-mn is 
produced at M2 in the Hypocysta-series, the crossveins are aligned in 
this tribe and in other members of the Ypthimini-section, resulting in 
a straight cell. The female foreleg is pentamerous in the present 
tribe, a situation which is never found in the coenonymphines and 
very rare in either the Maniolini or the Erebiini. 

MEM. AMER. ENT. SOC, 24 



82 



THE SATYRIDAE 



0.2 XML, FL 




Ypthima 

Thympia 

Xois 

Dallacha 

Kolasa 

Melampias 

Pseudonympha 

Strabena 

Callerebia 
Paralasa 
Loxeiebia 



Fig. 170. Satyrinae: Ypthimini. Relative lengths of the femur + tibia 
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) 
and hindlegs (HL, heavy stippling) of selected genera. In all instances the 
ML value is unity. The top bar for each genus represents the measurements 
obtained from males, the bottom bar those from females. 

All ypthimines are found in the Old World, and most genera are 
exclusively tropical. Most species fly in open country; few are found 
in scrub country, much less in the deep forest. Three major series 
are recognized in the Ypthimini, as follows: 

The Ypthima-senes, sensu stricto, characterized generally by their 
small size, brown coloration and diffuse androconia. This series is 
found throughout the Paleotropics. 



LEE D. MILLER 83 

The Callerebia-series, characterized by large size, brown color- 
ation and androconia arranged in rows on the discal portion of the 
forewing (Fig. 184). These species are generally distributed in the 
temperate mountains of the Middle and Far East. 

The Melampias-series, characterized generally by reddish shades 
on the upper surface, more elongate wings, small size and either 
diffuse androconia or none. These butterflies are found in South 
Africa and Madagascar. 

The Ypthimini are characterized as follows: 

The eyes are naked. The antennae are short, often less than 
two-fifths the length of the forewing costa (only one-fourth as long 
in Mashuna) and never as long as half the length of the costa (nearly 
half in Paralasa and Dallacha). The antennal club is variously 
developed: in such genera as Physcaeneura, Periplysia and Dallacha 
the club is less than twice as thick as the shaft, whereas in genera like 
Mashuna the club is almost four times as thick as the shaft. The 
third segment of the palpus is usually from one-fourth to slightly over 
one-third the length of the second, but the third segment is more 
than half as long as the second segment in Physcaeneura. The hairs 
of the second segment of the palpus are very long, usually three to 
five times as long as the segment is wide. 

The male foreleg is extremely reduced (Fig. 170), with the tarsus 
represented in most genera by a small knob on the end of the tibia 
(Fig. 173, for example). In such genera as Mashuna (Fig. 177), 
Strabena and Pandima, the tarsus is completely fused to the tibia, 
but in the Callerebia-series (Fig. 185) the tarsus is moderately well 
represented. The forefemur is as long as, or longer than, the fore- 
tibia. The female foreleg is also miniaturized, but it retains five 
tarsal subsegments, but the tarsus itself is significantly clubbed (Fig. 
174, for example). There are spines on the second, third and fourth 
subsegments. The midleg is subequal to the hindleg in the Callerebia- 
and Melampias-series, but significantly shorter in the Y pthima-series 
(Fig. 170). The midtibia is less than twice the length of the first 
midtarsal subsegment, is smooth dorsad, without a spine at the dorsal, 
distal end and with well to moderately well developed tibial spurs, 
except in Physcaenura (Fig. 179). 

The forewing cell is usually produced at the origin of Ms and 
not excavate (except in Periplysia, Fig. 180); the cell is about half 

MEM. AMER. ENT. SOC, 24 



84 



THE SATYRIDAE 




Figs. 171-180. Satyrinae: Ypthimini. 171. Ypthima philomela (Linne), 
S venation. 172. Y. philomela, palpus. 173. Y. philomela, 6 foreleg, in- 
cluding trochanter and distal portion of coxa. 174. Y. philomela, 9 foretarsus. 
175. Y. philomela, midleg. 176. Mashuna mashuna (Trimen), S venation. 

177. M. mashuna, $ foreleg, including trochanter and distal portion of coxa. 

178. Xois sesara Hewitson, venation of $ hindwing. 179. Physcaemira panda 
(Boisduval), midleg. 180. P. panda, ds venation. 

as long as the forewing costa. The forewing radial veins arise in one 
or two branches from the cell, and veins Rs and Mi are well separated 
at their origins. Vein M2 usually arises midway between Mi and M3, 
but nearer Mi in the Melampias-series; Cui arises nearer, or much 
nearer, Ms than CU2. Forewing vein Sc is always much thickened 
at its base, the cubital stem is generally inflated and 2A never is. 



LEE D. MILLER 85 

The hindwing cell is straight, longest at the origin of Mi and is 
about three-fifths the length of the wing measured to the end of Mn. 
Hindwing vein Sc + Ri is as long as, and more commonly longer 
than, 3A. Veins Ms and Cui arise separately, but often approximate, 
and M- arises somewhat nearer Mi than M3, or midway between the 
two veins. 

As noted in the general description, the pattern is rather constant. 
For illustrations of the various species see Moore ( 1893[1 890-1 893]) 
and van Son ( 1955 ). 

Genera Included in the Ypthimini 
Ypthima-sevies, sensu stricto 

Dallacha Moore, 1893(1890-1893): 94. Type-species: Yphthima (sic.) hya- 
griva Moore, by original designation. 

Hemadera Moore, 1893(1890-1893): 107. Type-species: Yphthima (sic.) 
narasingha Moore, by original designation. 

Kolasa Moore, 1893(1890-1893): 82. Type-species: Satyrus chenui Guerin, 
by original designation. 

Lohana Moore, 1893(1890-1893): 92. Type-species: Yphthima (sic.) inica 
Hewitson, by original designation. 

Mashuna van Son, (July) 1955: 159. Type-species: Ypthima mashuna Tri- 
men, by original designation. 

= Ypthimorpha Overlaet, (Dec.) 1955: 23. Type-species: Ypthima 
mashuna Trimen, by original designation. 

Nadiria Moore, 1893(1890-1893): 85. Type-species: Ypthima bolanica Mar- 
shall, by original designation. 

Pandima Moore, 1893(1890-1893): 86. Type-species: Satyrus nareda Kollar, 
by original designation. 

Thympia Moore, 1893(1890-1893): 58. Type-species: Papilio baldus Fabri- 
cius, by original designation. 

Xois Hewitson, 1865: 282. Type-species: Xois sesara Hewitson, by monotypy. 

Ypthima Hiibner, 1818(1818-1825): 17. Type-species: Papilio philomela 
Linne, by monotypy. This name is frequently misspelled in the litera- 
ture as "Yphthima". Hemming (1964c: 157-158) makes a strong case 
for "philomela" of Hiibner being a misidentification of Ypthima hueh- 
neri Kirby. Should the International Commission on Zoological No- 
menclature rule that this is the case, Y. huebncri will become the type- 
species. 

Ypthimomorpha van Son, 1955: 158. Type-species: Ypthima itonia Hewit- 
son, by original designation. 

MEM. AMER. ENT. SOC, 24 



86 



THE SATYRIDAE 




Figs. 181-188. Satyrinae: Ypthimini. 181. Melampias hyperbius (Lin- 
ne), $ venation. 182. Cassionympha cassius (Godart), 3 venation. 183. C. 
cassius, 9 foretarsus. 184. Callerebia scanda (Kollar), $ venation. 185. 
C. scanda, $ foreleg. 186. C. scanda, ? foretarsus. 187. C. scanda, mid- 
leg. 188. Boeberia parmenio (Boeber), $ venation. 



Melampias-series 

Callyphthima Butler, 1880: 335. 

by original designation. 
Cassionympha van Son, 1955: 96. 

original designation. 
Coenyropsis van Son, 1958: 6. Type-species: Satyrus natalii Boisduval, by 

original designation. 



Type-species: Psendonympha wardii Butler, 
Type-species: Satyrus cassias Godart, by 



LEE D. MILLER 87 

Neita van Son, 1955: 101. Type-species: Pseudonym pha neita Wallengren, 

by original designation. 
Melampias Hubner, [181 9] ( 1816-1826) : 63. Type-species: Papilio hyperbius 

Linne. designated by Scudder, 1875a: 214. 
Neocoenyra Butler, 1885: 758. Type-species: Neocoenyra duplex Butler, by 

monotypy. 
Periplysia Gerstacker, 1871: 358. Type-species: Periplysia leda Gerstacker, 

by monotypy. 
Pseudonym pha Wallengren, 1857: 31. Type-species: Papilio hippia Cramer, 

designated by Butler, 1868a: 194. 
Physcaeneura Wallengren, 1857: 32. Type-species: Satyrus panda Boisduval, 

by monotypy. 
Strabena Mabille, 1887: 10, 22. Type-species: Satyrus tamatavae Boisduval, 

by original designation. 
Stygionympha van Son, 1955: 137. Type-species: Pseudonympha vigilans 

Trimen, by original designation. 

Callerebia-series 

Argestina Riley, 1922: 469. Type-species: Callerebia waltoni Elwes, by origi- 
nal designation. 
Boeberia Prout, 1901: 233. Type-species: Papilio parmenio Boeber, by origi- 
nal designation. 

= Erebomorpha Elwes, 1899: 351. Type-species: Papilio parmenio 
Boeber, by original designation. Preoccupied by Erebomorpha 
Walker, 1860. 
Callerebia Butler, 1867e: 217. Type-species: Erebia scanda Kollar, by origi- 
nal designation. 
Loxerebia Watkins, 1925: 237. Type-species: Callerebia pratorum Oberthur, 

by original designation. 
Paralasa Moore, 1893(1890-1893): 103. Type-species: Erebia kalinda Moore, 
by original designation. 

Genus PAL AEON YMPHA Butler, 1871 
(Figures 189-195) 

Palaeonympha Butler, 1871: 401. Type-species: Palaeonympha opalina But- 
ler, by original designation. 

The original description of Palaeonympha pointed to its similarity 
to the North American Neonympha (Euptychiini). If Palaeonympha 
is a member of the Euptychiini it would be the first known Old World 
representative of that tribe. It is tempting to place the present genus 
there, since it is superficially very similar to the American Megisto 
cymela (Cramer), except for the androconial distribution in opalina. 

MEM. AMER. ENT. SOC, 24 



THE SATYRIDAE 



0.4X ML, FL 


ML 




HL 


IxmLi 


Y//////A 






fmlmmm 


?SMmM<M%&m 


1 




189 




i 



Palaeonympha 




Figs. 189-195. Satyrinae: triba uncertain, Palaeonympha opalina Butler. 
189. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diago- 
nal lines), midlegs (ML, light stippling) and hind'egs (HL, heavy stippling). 
The ML value is unity. The top bar represents the measurements obtained 
from the male, the bottom bar those from the female. 190. & venation. 191. 
antenna] club. 192. palpus. 193. 6 foreleg. 194. $ foretarsus. 195. midleg. 



There are certain similarities betwesn the Euptychiini and the Ypthi- 
mini — indeed, they probably arose from the same basic stock — and it 
is also tempting to place Palaeonympha in the latter tribe on purely 
geographical considerations, but if the present genus is one of the 
ypthimines, it is the only one with hairy eyes. Certainly Palaeo- 
nympha is either an ypthimine or a euptychiine, or it may represent 
an intermediate tribe, but I cannot place the genus in either tribe 
with certainty, and I hesitate to erect a monotypic tribe at this time. 
Palaeonympha is characterized as follows: 

The eyes are slightly hairy. The antennae are just less than 
half as long as the forewing costa. The antennal club is moderately 
well developed, occupying the distal quarter of the antenna and two 
and a half to three times as thick as the shaft. The third segment of 
the palpus is long, about two-fifths the length of the second. The 
hairs of the second segment of the palpus are three and a half times 
as long as the segment is wide. 



LEE D. MILLER 89 

The male foreleg is greatly reduced (Figs. 189, 193): the femur 
is longer than the tibia, and the tarsus is represented only by a bump 
at the end of the tibia. The female foreleg is also miniaturized (Figs. 
189, 194) with a pentamerous tarsus, the fifth subsegment of which 
is greatly reduced, bearing spines on the second and third subseg- 
ments. The hindlegs are slightly longer than the midlegs (Fig. 189). 
The midtibia is less than twice the length of the first midtarsal sub- 
segment, smooth dorsad, with no spine at the dorsal, distal end and 
with well developed tibial spurs. 

The forewing cell is very slightly excavate between Mi and M- 
and about half as long as the forewing costa. The forewing radial 
veins arise from the cell in a single branch, and Rs and Mi are 
separate at their origins. Vein M- arises slightly nearer Mi than Ma, 
and Cui arises nearer Ma than Cu-. Forewing vein Sc and the cubital 
stem are both much inflated basad, but 2A is not. 

The hindwing cell is somewhat produced at the origin of M3 and 
is about three-fifths the length of the wing measured to the end of 
Ms. Hindwing vein Sc + Ri is longer than 3 A, and Ma and Cui 
are well separated at their origins. Vein M 2 arises nearer Mi than Ma. 

The single species is brown above with a double-pupilled ocellus 
in Mi-M- of the forewing and single-pupilled ones in Rs-Mi and 
Ma-Cui of the hindwing. There is an androconial patch of mealy 
scales along the boundary of the cell and extending out the veins 
from Mi to 2A (Fig. 190). The under surface is light brown with 
discal and extradiscal transverse reddish-brown bands and well de- 
veloped ocelli as on the upper surface, plus very small ones in 
forewing spaces Ma-Cui through CU--2A and better developed ones 
in hindwing spaces M1-M2, Mi>-Ma and CU2-2A. 

Tribe Euptychiini, new tribe 
(Figures 196-210) 

This tribe shows the greatest affinities to the Old World Ypthi- 
mini and was probably derived from a common stock with it. 
The euptychiines are less closely related to the Maniolini and Coeno- 
nymphini. 

The Euptychiini are separable from the Ypthimini on the basis 
of the slightly longer and more square-cut forewing cell and the 
ocellus in forewng space Mi-Ma and differ from the coenonymphines 

MEM. AMER. ENT. SOC, 24 



90 



THE SATYRIDAE 



0.4XML 



IxML, 




Euptychia 

Taygetis 

Pindis 

Paramecera 

Oressinoma 



Fig. 196. Satyrinae: Euptychiini. Relative lengths of the femur + 
tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip- 
pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances 
the ML value is unity. The top bar for each genus represents the measure- 
ments obtained from males, the bottom bar those from females. 

in that the latter have an excavate forewing cell and a greater abor- 
tion of the female foreleg, as well as a greater distention of forewing 
vein 2A. 

All of the genera definitely assigned to the euptychiines are found 
in the New World, from southern Canada through the Neotropics to 
temperate South America. Most are woods or forest dwellers, in 
contrast to most members of the Ypthimini-section. The most com- 
prehensive review of the tribe to date is included in Forster ( 1964), 
but, unfortunately, it is by no means complete. The Euptychiini 
are characterized below. 

The eyes are variously naked or hairy. The antennae are short, 
just under to just over two-fifths the length of the forewing costa. 
The antennal club is usually weakly developed and about twice the 
thickness of the shaft (about three times as thick in Cyllopsis). The 
third segment of the palpus is about a third as long as the second 
segment. The hairs of the second segment of the palpus are variable: 
in the tropical groups they are about three times as long as the 
segment is wide, but in the Nearctic Me gist o cymela (Cramer) the 
hairs are almost five times the width of the segment. 

The male foreleg is reduced, with a monomerous, unspined tarsus; 
the femur and tibia are of about the same length. The female foreleg 



LEE D. MILLER 



91 




Figs. 197-205. Satyrinae: Euptychiini. 197. Euptychia mollina Hiibner, 
8 venation. 198. E. mollina, palpus. 199. E. mollina, $ foreleg. 200. E. 
mollina, 2 foretarsus. 201. Cissia penelope (Fabricius). 9 foretarsus. 202. 
E. mollina, midleg. 203. Megisto cymela (Cramer), midleg. 204. Cyllopsis 
hedemanni R. Felder, S venation. 205. Taygetis mermeria (Cramer), S 
venation. 

is also reduced, with a pentamerous, clubbed tarsus bearing spines 
on the second, third and fourth subsegments (also on the first in at 
least Neonympha). The midtibia is one and a half to one and three- 
quarters times the length of the proximal midtarsal subsegment. The 
midtibia is variously smooth dorsad in most genera to slightly spiny 
in such genera as Neonympha and spiny in such genera as Cissia. 
The tibial spurs are present and well developed in most species, but 

MEM. AMER. ENT. SOC, 24 



92 THE SATYRIDAE 

absent in Euptychia mollina Hiibner, and there is no spine at the 
dorsal, distal end of the midtibia. The relative lengths of the legs 
of selected Euptychiini are shown in Fig. 196. 

The forewing cell is generally square-cut and varies little from 
half the length of the forewing costa. The forewing radial veins 
arise from the cell in two branches (rarely one branch), and veins 
Rs and Mi arise well separate. Vein M2 arises somewhat nearer Mi 
than M a , and Cui arises nearer M 3 than CU2. Forewing veins Sc and 
the cubital stem are basally inflated in all genera, and 2A is inflated in 
such genera as Taygetis and Oressinoma (Figs. 205, 210, respec- 
tively). The latter genus displays an unusual form of inflation of 
the veins. 

The hindwing cell is straight and produced at the origin of M3; 
the cell is half to three-fifths the length of the wing measured to the 
end of Mm. Hindwing veins Sc + Ri and 3 A are of about the same 
length, Ms and Cui arising well separate and M2 arising nearer, 
occasionally much nearer, Mi than M3. 

The pattern is reasonably constant. Most species are brown 
above — they may be white or iridescent purple, blue in part — with 
or without ocelli. The under surface is usually brown with trans- 
verse darker markings and a more or less complete complement of 
ocelli. Oressinoma typhla Westwood is aberrant: white above and 
below bordered with blackish brown and with reddish lunules in the 
dark border. 

Genera Included in the Euptychiini 1 

'Names preceded by asterisks (*) are genera proposed too late for de- 
tailed study, but from the original descriptions they are judged to be members 
of this tribe. 

*Archeuptychia Forster, 1964: 80. Type-species: Papilio c'uena Drury, by 
original designation. 

*Caerul euptychia Forster, 1964: 92. Type-species: Euptychia caerulea But- 
ler, by original designation. 

*Capronnieria Forster. 1964: 73. Type-species: Euptychia abretia Capron- 
nier, by original designation. 

*Cepheuptychia Forster, 1964: 96. Type-species: Euptychia cephus Butler, 
by original designation. 

*Chlor euptychia Forster, 1964: 119. Type-species: Papilio chloris Cramer,, 
by original designation. 

Cissia Doubleday, 1848b: 33. Type species: Papilio clarissa Cramer (=Pa- 



LEE D. MILLER 



93 




Figs. 206-210. Satyrinae: Euptychiini. 206. Paramecera xicaque (Rea- 
kirt), 6 venation. 207. P. xicaque, palpus. 208. P. xicaque, £ foreleg. 209. 
P. xicaque, 2 foretarsus. 210. Oressinoma typhla Westwood, 6 venation. 

pilio penelope Fabricius), by monotypy. 

= Argyreuptychia Forster, 1964: 123. Type-species: Papilio penel- 
ope Fabricius, by original designation. 

*Coeruleotaygetis Forster, 1964: 73. Type-species: Euptychia periboea God- 
man and Salvin, by original designation. 

Cyllopsis R. Felder, 1869: 474. Type-species: Cyllopsis hedemanni R. Felder, 
by monotypy. 

*Erichthodes Forster, 1964: 118. Type-species: Euptychia erichtho Butler, 
by original designation. 

Euptychia Hlibner. 1818 (1818-1825): 20. Type-species: Euptychia mol- 
lina Hubner, designated by Hemming, 1937: 150. Butler (1868a: 
194) designated Papilio herse Cramer as the type, based on Euptychia 
Hubner, [1819]( 1816-1826) : 54, but this designation was invalidated 
when the appropriate portions of the "Zutrage" were found to be older 
than those of the "Verzeichniss". 

*Euptychoides Forster, 1964: 97. Type species: Euptychia saturnus Butler, 
by original designation. 

*Godartiana Forster, 1964: 118. Type-species: Satyrus byses Godart, by 
original designation. 

*Harjesia Forster, 1964: 78. Type-species: Taygetis blanda Moschler, by 
original designation. 

*Haywardina Forster, 1964: 109. Type-species: Satyrus necys Godart, by 
original designation. Preoccupied by Haywardina Aczel, 1952, but no 
replacement name is proposed. 

*Hermeuptychia Forster, 1964: 87. Type-species: Papilio hermes Fabricius, 
by original designation. 



MEM. AMER. ENT. SOC, 24 



94 THE SATYRIDAE 

*Magneuptychia Forster, 1964: 125. Type-species: Popilio libye Linne, by 

original designation. 
*Megeuptychia Forster, 1964: 122. Type-species: Nymphalis autonoe Cramer, 

by original designation. 
Megisto Hiibner, [1819]( 1816-1 826) : 54. Type-species: Papilio eurytus Fa- 

bricius (=PapiIio cymela Cramer), designated by Butler, 1868b: 14. 
*Moneuptychia Forster, 1964: 92. Type-species: Euptychia soter Butler, by 

original designation. 
Neonympha Hiibner, 1818(1818-1825): 8. Type-species: Papilio areolatus 

Smith and Abbot, designated by Hemming, 1937: 150. 
Oressinoma Westwood, 1852, in Doubleday, Westwood and Hewitson, 1846- 

1852: 371. Type-species: Oressinoma typhla Westwood, by monotypy. 

= Ocalis Westwood, 1852, in Doubleday, Westwood and Hewitson, 

1846-1852: 371. Type-species: Oressinoma typhla Westwood, by 

monotypy. Listed in synonymy of Oressinoma Westwood. 

Paramecera Butler, 1868b: 98. Type-species: Neonympha xicaque Reakirt, 

by monotypy. 
*Parataygetis Forster, 1964: 79. Type-species: Taygetis albinotata Butler, by 

original designation. 
*Parenptychia Forster, 1964: 83. Type-species: Papilio hesione Sulzer, by 

original designation. 
* Parypththimoid.es Forster, 1964: 106. Type-species: Neonympha poltys 

Prittwitz, by original designation. 
*Pharneuptychia Forster, 1964: 91. Type-species: Satyrus phares Godart, by 

original designation. 
Pindis R. Felder, 1869: 475. Type-species: Pindis squamistriga R. Felder, by 

monotypy. 
*Posteuptychia Forster, 1964: 137. Type-species: Euptychia mycalesoides 

Felder, by original designation. 
*Posttaygetis Forster, 1964: 74. Type-species: Papilio penelea Cramer, by 

original designation. 
*Praefaunula Forster, 1964: 137. Type-species: Euptychia armilla Butler, by 

original designation. 
*Pseudeuptychia Forster, 1964: 86. Type-species: Euptychia languida Butler, 

by original designation. 
*Pseudodebis Forster, 1964: 75. Type-species: Papilio valentina Cramer, 

by original designation. 
*Rareuptychia Forster, 1964: 87. Type-species: Euptychia clio Weymer, by 

original designation. 
*Satyrotaygetis Forster, 1964: 73. Type-species: Euptychia satyrina Bates, 

by original designation. 
*Splendeuptychia Forster, 1964: 128. Type-species: Euptychia ashna Hew- 
itson, by original designation. 
*Taygetina Forster, 1964: 77. Type-species: Taygetis hanghaasi Weymer, by 

original designation. 



LEE D. MILLER 95 

Taygetis Hiibner, [1819]( 1816-1826) : 55. Type-species: Papilio mermeria 
Cramer, designated by Westwood, 1852, in Doubleday, Westwood and 
Hewitson, 1846-1852: 355. Hemming (in press) accepts the designa- 
tion of Papilio virgilia Cramer as the type-species by Butler, 1868a: 194, 
but the intent of Westwood to designate mermeria as the type-species 
is unmistakable. 

*Vareuptychia Forster, 1964: 125. Type-species: Euptychia similis Butler, 
by original designation. 

*Weymerana Forster, 1964: 108. Type-species: Euptychia viridicans Wey- 
mer, by original designation. 

*Ypthimoides Forster, 1964: 100. Type-species: Neonympha yphthima Felder, 
by original designation. 

*Zischkaia Forster, 1964: 116. Type-species: Euptychia jumata Butler, by 
original designation. 

Tribe Coenonymphini, new tribe 
(Figures 211-222) 

The Coenonymphini are intimately related to the ypthimines 
and probably replace them in the north temperate zone savannas of 
both hemispheres. There are some relationships suggested with the 
maniolines, the latter group occupying scrub country environments: 
probably both evolved from a basic ypthimine stock. There is little 
to connect the coenonymphines with the Euptychiini or the Erebiini, 
two isolated tribes of the Ypthimini-section. 

The excavate forewing cell serves to distinguish most coeno- 
nymphines from most ypthimines, and the very long midtibia (over 
twice as long as the first midtarsal segment in all genera but Aphan- 
topus) is unique in the Ypthimini-section. The female foreleg of the 
Ypthimini and the Euptychiini is fully formed, having five tarsal sub- 
segments, whereas that of the Coenonymphini is lacking at least one 
of the subsegments. 

The Coenonymphini are exclusively Holarctic and are grassland 
species. Almost all are temperate, few (e. g., Coenonympha mix- 
turata Alpheraky) being found so far north as the subarctic. The 
Coenonymphini are characterized as follows: 

The eyes are naked. The antennae are always shorter than half 
the length of the forewing costa, and those of Lyela and Triphysa 
are less than two-fifths the length of the forewing. The antennal club 
is variable in its development: in Aphantopus the club is only about 

MEM. AMER. ENT. SOC, 24 



96 



THE SATYRIDAE 



0.2*ML 




Fig. 211. Satynnae: Coenonymphini. Relative lengths of the femur + 
tibia + tarsus of the forelegs (FL diagonal lines), midlegs (ML, light stip- 
pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances 
the ML value is unity. The top bar for each genus represents the measure- 
ments obtained from males, the bottom bar those from females. 

two and a half times as thick as the shaft, five times as thick in Tri- 
physa (Fig. 220) and seven times the thickness of the shaft in Lyela. 
The third segment of the palpus is generally half to one-third as long 
as the second, but the third segment is less than one-sixth the length 
of the second in Triphysa and Lyela. The hairs of the second seg- 
ment of the palpus are about four times as long as the segment is 
wide. 

The male foreleg is weakly developed with a monomerous, un- 
spined (weakly spined in Lyela) tarsus. The forefemur and fore- 
tibia are of about the same length; in some genera slight variations 
may be apparent. The female foreleg is reduced and is of two gen- 
eral types: in such genera as Coenonympha there are four tarsal sub- 
segments which are spined on at least the third and fourth ones, 
whereas in other genera like Chortobius, Lyela and Triphysa there 
is but one, unspined tarsal subsegment. With the exception of 
Aphantopus, the midtibia is very long, greater than twice as long as 
the first midtarsal subsegment. The tibial spurs are present and well 
developed in all genera but Triphysa, in which they are totally absent. 
The midtibia is smooth dorsad and has no spine at the dorsal, distal 
end. The hindlegs are significantly longer than the midlegs (Fig. 
211). 

The forewing cell is deeply excavate and varies little from half 
the length of the forewing costa. The forewing radial veins generally 
arise in two branches from the cell, but in three branches in Aphan- 



LEE D. MILLER 



97 



topus; veins Rs and Mi are separate at their origins, although they 
may be approximate. Vein M- arises nearer Mi than M :! , and Cui 
arises midway between Ms and Cu^, or slightly nearer M3. Fore- 




Figs. 212-222. Satyrinae: Coenonymphini. 212. Coenonympha oedippus 
(Fabricius). 6 venation. 213. C. oedippus, antennal club. 214. C. oedippus, 
palpus. 215. C. oedippus, 6 foreleg. 216. Chortobius pamphilus (Fabricius). 
9 foretarsus. 217. C. oedippus, midleg. 218. C. oedippus, 9 foretarsus. 219. 
Triphysa phryne (Pallas), 6 venation. 220. T. phryne, antennal club. 221. 
T. phryne, midleg. 222. Aphantopus hyperantus (Linne), 6 venation. 

wing veins Sc, the cubital stem and 2A are all inflated, though the 
latter may be only slightly thickened. 

The hindwing cell is square-cut distad and half to three-fifths 
the length of the wing to the end of M3. Hindwing vein Sc + Ri is 

MEM. AMER. ENT. SOC, 24 



98 THE SATYRIDAE 

as long as, or longer than, 3A. Veins Ms and Cui are separate, 
but close together, at their origins, and M2 arises nearer, or much 
nearer, Mi than Ma. 

The pattern is fairly constant. The upper surface is brown to 
tawny with few, if any, ocelli. The under surface of the forewings 
is likewise brown to tawny, and the hindwings are brown to grayish- 
olive, often with a light mesial band, with a more or less developed 
series of ocelli on both wings. Triphysa phryne (Pallas) is the only 
species to show marked sexual dimorphism: the male is brown with 
a more complete set of ocelli than is usual in the tribe, and the female 
is white with a full complement of ocelli. 

Genera Included in the Coenonymphini 

Aphantopus Wallengren, 1853: 30. Type-species: Papilio hyperantus (mis- 
spelled "hyperanthus") Linne, by monotypy. 

Chortobius [Dunning and Pickard], 1858: 5. Type-species: Papilio pamphilus 
Linne, designated by Moore, 1893 (1890-1893): 51-52. 

Coenonympha Hiibner, [18 19] ( 1816-1826) : 65. Type-species: Coenonympha 
oedipe Hiibner (=Papilio oedippus Fabricius), designated by Butler, 
1868a: 194 (as Papilio geticus Esper). 

Dubierebia Muschamp, 1915b: 20. Type-species: Coenonympha myops 
Staudinger, by original designation. 

Lyela Swinhoe, 1908: 60. Type-species: Lyela macmahoni Swinhoe, by origi- 
nal designation. 

Sicca Verity, 1953: 83. Type-species: Papilio dorus Esper, by original desig- 
nation. Proposed as a subgenus of Coenonympha Hiibner. 

Triphysa Zeller, 1850: 308. Type-species: Papilio tircis Stoll (= Papilio 
phryne Pallas), designated by Butler, 1868a: 194. 

=Phryne Herrich-Schaffer, 1844 (1843-1856): 90. Type-species: 
Papilio tircis Stoll (=Papilio phryne Pallas), by monotypy. Pre- 
occupied by Phryne Meigen, 1 800. 

Tribe Maniolini Hampson, 1918 

(Figures 223-232) 

Maniolinae Hampson, 1918: 385; Maniolidi Verity, 1953: 228, 237. 

This tribe is most closely related to the Coenonymphini; probably 
both tribes evolved in the Holarctic from a basic ypthimine stock. 
The maniolines are much more distantly related to either the Eupty- 
chiini or the Erebiini. 

The maniolines are distinguished from other members of the 



LEE D. MILLER 



99 



Ypthimini-section by forewing vein M-2 arising midway between Mi 
and M.i (in other tribes M2 arises nearer Mi), and by the distribu- 
tion of androconia. Whereas the Euptychiini and the Ypthimini 



0.2 x ML, FL 


ML 


HL IxML, 


1 






1 












r 










v//A 










1 








1 



Maniola 



Cercyonis 



Fig. 223. Satyrinae: Maniolini. Relative lengths of the femur + tibia 
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and 
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML 
value is unity. The top bar for each genus represents the measurements 
obtained from males, the bottom bar those from females. 

have a pentamerous female foreleg, this tribe shows leg abortion in 
all genera, like the Coenonymphini. 

Cercyonis is considered by many authors (e.g., Hemming, in 
press ) to be biologically synonymous with Minois Hiibner. The latter 
genus, however, is a member of the Satyrini, as shown by the pro- 
duction of the hindwing cell along vein Mi> and the typically satyrine 
configuration of the midleg (Fig. 301). The midleg of Cercyonis 
is typical of the Maniolini (Fig. 228) and has nothing in common 
evolutionary with Minois. The proper assignment of Cercyonis 
again demonstrates the necessity of looking beyond mere superficial 
similarities. 

The maniolines are found throughout the north temperate zone, 
and a few members are also found in the Arctic. These butterflies 
seem to replace the grassland Coenonymphini in scrublands. The 
Maniolini are characterized as follows: 

The eyes are naked. The antennae are greater than two-fifths, 
but less than half, the length of the forewing costa. The antennal club 
is weakly developed, from less than twice to about two and a half 
times the thickness of the shaft, and occupying the distal third to a 
quarter of the antenna. The third segment of the palpus is about 
one-third of the length of the second. The hairs of the second seg- 
ment of the palpus are long, about four to five times as long as the 
segment is thick. 

The male foreleg is greatly reduced, the femur and the tibia are 

MEM. AMER. ENT. SOC, 24 



100 



THE SATYRIDAE 



of approximately the same length and the tarsus is monomerous and 
unspined. The female foreleg is also greatly reduced with a mono- 
or dimerous tarsus without spines. The mid- and hindlegs are of 




Figs. 224-232. Satyrinae: Maniolini. 224. Maniola jurtina (Linne), 
$ venation. 225. M. jurtina, palpus. 226. M. jurtina, $ foreleg. 227. M. 
jurtina, 9 foretarsus. 228. M. jurtina, midleg. 229. Pyronia tithonus (Lin- 
ne), $ venation. 230. Cercyonis pegala (Fabricius), 6 venation. 231. C. 
pegala, S foreleg. 232. C. pegala, 2 foretarsus. 

about the same length (Fig. 223). The midtibia is somewhat less 

than twice as long as the basal midtarsal subsegment, without spines 

dorsad or at the dorsal distal end and with well developed tibial spurs. 

The forewing cell is somewhat excavate and rather uniformly half 



LEE D. MILLER 101 

as long as the forewing costa. The forewing radial veins arise in 
three branches from the cell, and veins Rs and Mi are separate at 
their origins. Vein M- arises about midway between Mi and M:s. 
and Cui arises nearer Ma than Cu-. Forewing vein Sc is always 
inflated basally; the cubital stem and 2A may be greatly inflated 
(Maniola, Fig. 224) or not noticeably inflated (Cercyonis, Fig. 230). 

The hindwing cell is produced at the origin of Ma and varies from 
less than half to as long as three-fifths the length of the wing measured 
to the end of Ma. Vein Sc + Ri is longer than 3 A, and Ma and Cui 
are well separated at their origins. Vein M- arises nearer Mi than Ma. 

The upper surface is either tawny with brown margins or brown 
with or without a yellow discal patch on the forewing with the ocelli, 
if they are present, in the forewing spaces Mi-Ma and Cua-2A and on 
the hindwing at the tornus. Beneath the wings are light brown with 
darker brown striae and moderately well developed ocelli on both 
wings. 

Genera Included in the Maniolini 

Cercyonis Scudder, 1875b: 241. Type-species: Papilio alope Fabricius ( = 

Papilio pegala Fabricius), by original designation. 
Hyponephele Muschamp, 1915a: 156. Type-species: Papilio lycaon Rottem- 

burg, by monotypy. 
I data deLesse. 1952: 72. Type-species: Pyronia cecilia Vallantin. by original 

designation. Proposed as a subgenus of Pyronia Hiibner. 
Maniola Schrank, 1801: 152. Type-species: Maniola lemur Schrank (=Papilio 
jurtina Linne), designated by Scudder. 1875a: 211. 

= Epinephele Hiibner, [1819]( 1816-1826) : 59. Type-species: Pa- 
pilio janira Linne (= Papilio jurtina Linne), designated by Butler, 
1868a: 194. Note that many authors have misspelled this name 
as "Epinephile" or "Epinephild". 
Pasiphana de Lesse, 1952: 72. Type-species: Hipparchia bathseba Fabricius, 
by original designation. Proposed as a subgenus of Pyronia Hiibner. 
Pyronia Hiibner, [ 1 8 1 9] < 1816-1826) : 59. Type-species: Pyronia tithone 
Hiibner (=Papilio tithonus Linne), designated by Scudder, 1875a: 261. 

Tribe Erebiini Tutt, 1896 

(Figures 233-246) 
Erebiinae Tutt, 1896: 87. 402. 

This tribe shows certain affinities with the Ypthimini and to the 
Satyrini-section, but is closer to the former; the similarity to the latter 

MEM. AMER. ENT. SOC, 24 



102 



THE SATYRIDAE 



is probably convergence. Structurally no group is exceptionally close 
to the erebiines, except the Callerebia-series of the Ypthimini. 



0.2*ML FL 




Erebia 

Medusia 

Phorcis 

Gorgo 

Atercoloratus 



Fig. 233. Satyrinae: Erebiini. Relative lengths of the femur + tibia 
+ tarsus of the forelegs (FL, diagonal lines), midlegs (ML. light stippling) and 
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML 
value is unity. The top bar for each genus represents the measurements 
obtained from males, the bottom bar those from females. 

The erebiines may be distinguished from other satyrines of the 
Ypthimini-section by the extremely short third palpal segment, which 
is less than a fourth as long as the second segment in all genera. The 
facies of the species are characteristic, but, as indicated in the dis- 
cussions of the Dirini, the Hypocystini and the Pronophilini, some 
other genera outside the present tribe share the dark coloration, 
rounded wings and alpine habits with these butterflies. The very 
slight inflation of the forewing veins, even of Sc, is atypical of the 
Ypthimini-section. 

All of the erebiines are Holarctic, and many species are found 
in the high Arctic, as well as in alpine situations; few are at home in 
truly temperate climates. Warren (1936) has admirably mono- 
graphed this group. The Erebiini are characterized as follows: 

The eyes are naked. The antennae are from just over two-fifths 
to greater than half the length of the forewing costa. The antennal 
club is moderately well to very well developed, generally occupying 
the distal quarter of the antenna and inflated as much as five times 
the thickness of the shaft. The third segment of the palpus is very 
short, from one-quarter to one-sixth the length of the second. The 



LEE D. MILLER 



103 




Figs. 234-246. Satyrinae: Erebiini. 234. Erebia ligea (Linne), 6 vena- 
tion. 235. E. ligea, antennal club. 236. E. ligea. palpus. 237. E. ligea, 8 
foreleg. 238. E. ligea, 9 foretarsus. 239. E. ligea, midleg. 240. Phorcis 
epistygne Hiibner, <5 venation. 241. P. epistygne, S foreleg. 242. P. epi- 
stygne, 9 foretarsus. 243. P. epistygne, midleg. 244. Atercoloratus alini 
(Bang-Haas), 6 venation. 245. A. alini, palpus. 246. A. alini, $ foreleg, in- 
cluding trochanter and distal portion of coxa. 

MEM. AMER. ENT. SOC, 24 



104 THE SATYRIDAE 

hairs on the second segment of the palpus are long, between three 
and five times as long as the segment is wide. 

The male foreleg is reduced with a mono- or dimerous, unspined 
or weakly spined tarsus; the femur is generally as long as, or longer 
than, the tibia. The development of the female foreleg is variable, 
though the leg itself is always greatly miniaturized (Fig. 233). 
There are two to five tarsal subsegments with spines on all but the 
last one. The midleg is about as long as, and frequently longer 
than, the hindleg (Fig. 233). The midtibia is from just under to 
well over one and a half times the length of the first midtarsal sub- 
segment. The midtibia never bears a spine at its dorsal, distal end 
and is usually smooth dorsad, but slightly spiny in Phorcis (Fig. 243). 
The tibial spurs are present and well developed. 

The forewing cell is slightly excavate and varies little from half 
the length of the forewing costa. The forewing radial veins arise in 
three branches from the cell, and veins Rs and Mi are well separated 
at their origins. Vein M_> arises much nearer Mi than Ms, and Cui 
arises nearer M3 than Cu^>. Forewing vein Sc is always more or less 
inflated, the cubital stem is somewhat inflated and 2A hardly at all. 

The hindwing cell is produced at the origin of M3 and is half to 
three-fifths as long as the wing measured to the end of Ms. Vein 
Sc + Ri is longer than 3 A, and M3 and Cui are well separated at 
their origins. Vein M2 arises nearer Mi than Ms. 

These are dark insects with very little pattern on either surface 
except scattered ocelli and a forewing tawny to russet shad 2; either or 
both pattern features may be absent. Illustrations of the species are 
given by Warren (1936). 

Genera Included in the Erebiini 

Atercoloratus Bang-Haas, 1938: 178. Type-species: Coenonympha alini Bang- 
Haas, by original designation. 

Erebia Dalman, 1816: 58. Type-species: Papilio ligea Linne, by original 
designation. 

= Epigea Hiibner, [1819]( 1816-1826) : 62. Type-species: Papilio 
ligea Linne, designated by Hemming, 1933: 198. 

Gorgo Hiibner, [18 19] ( 1816-1826 ) : 64. Type-species: Papilio ceto Hiibner, 
designated by Hemming. 1933: 198. 

Marica Hiibner, [1 8 19] ( 1816-1826) : 63. Type-species: Papilio stygne Ochs- 
enheimer, designated by Hemming, 1933: 198. 

Medusia Verity, 1953: 179. Type-species: Papilio medus Schiffermiller, by 
original designation. Proposed as a subgenus of Erebia Dalman. 



LEE D. MILLER 105 

Oreina Westwood, 1841: 76. Type-species: Papilio cassiope Fabricius, desig- 
nated by Butler, 1868a: 194. 

Phorcis Hubner, [ 18 19]( 181 6-1826) : 62. Type-species: Phorcis epistygne 
Hiibner, designated by Hemming, 1933: 198. 

Simplica Verity, 1953: 194. Type-species: Papilio epiphron Knoch, by origi- 
nal designation. Proposed as a subgenus of Erebia Dalman. 

Syngea Hubner, [18 1 9] ( 1816-1826) : 62. Type-species: Papilio pronoe Esper, 
designated by Hemming, 1933: 198. 

Triariia Verity. 1953: 186. Type-species: Papilio triarius de Prunner, by 
original designation. Proposed as a subgenus of Erebia Dalman. 

Truncaefalcia Verity, 1953: 188. Type-species: Papilio aethiops Esper, by 
originial designation. Proposed as a subgenus of Erebia Dalman. 

Tribe Dirini, new tribe 
(Figures 247-259) 

This tribe is not as primitive as the hypocystines, as may be seen 
form examination of the forelegs of both sexes, even though it seems 
to connect the remainder of the Satyrini-section with the South Afri- 
can members of the Lethini — a convergence which is probably 
ecological and with little phylogenetic significance. The forelegs of 
both sexes are greatly reduced, about as much so as in any tribe; 
the dirines are nevertheless primitive members of the Satyrini-section. 
There is little to connect the Dirini — or any member of the Satyrini- 
section, for that matter — with the Hypocystini except the configura- 
tion of the hindwing cell (Figs. 248, 254, 257). In common with 
the Satyrini, Pronophilini (usually to a lesser extent) and Melanar- 
giini are the greatly reduced forelegs, the spiny midtibiae and the well 
developed tibial spurs of the present tribe. Some species have a well 
developed dorsal, distal spine at the end of the midtibia (Fig. 259), 
a characteristic shared with the Satyrini and indicating a close rela- 
tionship between these two tribes. 

The Dirini are characterized chiefly by the short forewing cell, 
always less than half the length of the wing; in other Satyrinae the 
cell is half the length of the wing or greater. The forewing cell is 
also rounded and reminiscent of that of the Lethini. 

All members of this tribe are restricted to South Africa and are 
well illustrated by van Son (1955). The Dirini are characterized 
as follows: 

The eyes are hairy in all genera. The antennae are approximately 
two-fifths the length of the forewing costa. The antennal club is 

MEM. AMER. ENT. SOC, 24 



106 



THE SATYRIDAE 



0.2XML, FL 




Dira 



Tarsocera 



Torynesis 



Fig. 247. Satyrinae: Dirini. Relative lengths of the femur + tibia + 
tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and 
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML 
value is unity. The top bar for each genus represents the measurements ob- 
tained from males, the bottom bar those from females. 

abrupt, occupying the distal one-fifth to one-third of the antenna 
and three to five times as thick as the shaft (Fig. 249). The third 
segment of the palpus is usually about a third the length of the 
second, but is half as long in Cassus (Fig. 254) and only one-ninth 
as long in Torynesis (Fig. 257). The hairs of the second palpal 
segment are two or three times as long as the segment is wide. 

The male foreleg is greatly reduced (Fig. 247), particularly as 
regards the tibia and tarsus. The femur is always more than twice 
as long as the tibia, and the tarsus has one or two unspined sub- 
segments (Figs. 251, 258). The female foreleg is also greatly re- 
duced, frequently as much so as that of the male, with a mono- or 
dimerous, unspined tarsus (Fig. 252). The midleg is as long as, 
and occasionally much longer than (Torynesis), the hindleg (Fig. 
247). The midtibia is almost twice as long as the first midtarsal 
subsegment in some genera such as Dira (Fig. 253), but in Tory- 
nesis (Fig. 259) and Dingana the midtibia is only slightly longer 
than is the proximal midtarsal subsegment. The midtibia is spiny 
dorsad and bears well developed tibial spurs; in most genera there is 
no spine at the dorsal, distal end of the midtibia, but the spine is 
weakly developed in Dingana. The configuration of the midleg in 
Dingana and Torynesis strongly suggests that of the Satyrini. 

The forewing cell is generally rounded distad and is greater than 
two-fifths, but less than half, the length of the forewing costa. The 
forewing radial veins arise from the cell in three branches, and Rs 
and Mi are well separated at their origins. Vein M2 arises nearer Mi 
than M3, and Cui arises nearer M3 than Cu_>. Only forewing vein 



LEE D. MILLER 



107 




Figs. 248-259. Satyrinae: Dirini. 248. Dira clytus (Linne), S venation. 
249. D. clytus, antennal club. 250. D. clytus, palpus. 251. D. clytus, $ fore- 
leg. 252. D. clytus, $ foretarsus. 253. D. clytus, midleg. 254. Cassus cassus 
(Linne), palpus. 255. C. cassus, S venation. 256. Torynesis mintha (Geyer), 
$ venation. 257. T. mintha, palpus. 258. T. mintha, $ foreleg. 259. T. 
mintha, midleg. 



Sc is inflated basad, and it is only slightly thickened. 

The hindwing cell is somewhat produced at the origin of M3 
and about half the length of the wing measured to the end of Ms. 
Hindwing veins Sc + Ri and 3 A are of about the same length, and 
veins Ms and Cui arise separately, though they may be approximate. 
Vein Mi» arises much nearer Mi than M3. 

MEM. AMER. ENT. SOC, 24 



108 THE SATYRIDAE 

These butterflies are dark colored insects on the upper surface 
with ocelli in forewing spaces M1-M3 and Qi2-2A. On the under 
surface the forewing ocelli are repeated and the hind wings are pat- 
terned by pale mottling and veinal markings. The general aspect 
is that of the Erebiini or the Satyrini. The species are figured by 
van Son (1955). 

Genera Included in the Dirini 

Cassus van Son, 1955: 79. Type-species: Papilio cassus Linne, by original 

designation. 
Dingana van Son, 1955: 70. Type-species: Leptoneura dingana Trimen, by 

original designation. 
Dira Hiibner, [ 1 819]( 1816-1 826) : 60. Type-species: Dira clyte Hiibner ( = 
Papilio clytus Linne), designated by Scudder, 1875a: 157. 

= Leptoneura Wallengren, 1857: 31. Type-species: Papilio clytus 
Linne, by monotypy. 
Tarsocera Butler, 1899: 903. Type-species: Leptoneura cassina Butler, by 

monotypy. 
Torynesis Butler, 1899: 903. Type-species: Dira mintha Geyer, by monotypy. 
=Mintha van Son, 1955: 76. Type-species: Dira mintha Geyer, by 
original designation. 

Tribe Pronophilini Clark, 1947 
(Figures 260-294) 
Pronophilinae Clark, 1947: 149. 

This tribe was derived from a basic Satyrini-like stock of Old 
World origin but is more primitive than any present-day Satyrini, 
particularly as regards the forelegs, hence the pronophilines are placed 
before the satyrines in this discussion. In addition to the affinity with 
the Satyrini, the Pronophilini share some characteristics (though not 
particularly important ones) with the South African Dirini, which 
are, in turn, the most primitive tribe within the Satyrini-section. The 
pronophilines are only very distantly related to the .A>n/ca-series of 
the Hypocystini; hence, any trans- Antarctic distribution of the family 
is doubtful. 

As in the Hypocystini and Dirini, convergent evolution toward 
the Holarctic Erebia is shown by several high-altitude pronophilines 
of the Lymanopoda-series. Several members of the Elina-series of. 
the present tribe show resemblance to members of the Holarctic 



LEE D. MILLER 



109 



0.4XML, FL 




Pronophi la 



Daedalma 



Eteona 



Steroma 



Gyrocheilus 



Elina 



Argyrophorus 



Neomaenas 



Nelia 



Lymanopoda 



Homoeonympha 



Sarromia 



Fig. 260. Satyrinae: Pronophilini. Relative lengths of the femur + 
tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip- 
pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances 
the ML value is unity. The top bar for each genus represents the measure- 
ments obtained from males, the bottom bar those from females. 

Maniolini; indeed, they were long considered members of Maniola 
( — Epinephele ) , but on structural grounds these butterflies are per- 
fectly valid pronophilines. Many aberrant patterns have evolved in 
the great adaptive radiation of this tribe in the Neotropics: the Chilean 



MEM. AME1 FNT. SOC, 24 



110 THE SATYRIDAE 

species Argyrophorous argenteus Blanchard is silver above, while 
Lymanopoda samius Westwood is caerulean blue on the upper surface 
and members of Drucina resemble nymphalids or danaids more than 
they do other satyrids. 

The Pronophilini are the only Neotropical satyrids which have 
the hindwing crossvein mi-ni2 produced distad at M 2 and hindwing 
vein Sc + Ri completely fused throughout its entire length. The 
present tribe can be distinguished from the Satyrini by the longer 
midtibia — never less than one and a half times the length of the first 
midtarsal subsegment — and the lack of a dorsal, distal midtibial spine. 
The more aborted forelegs serve to distinguish these butterflies from 
the Hypocystini. 

All members of the Pronophilini are Neotropical with the excep- 
tion of a single genus (Gyrocheilus) which penetrates the southern- 
most Nearctic. Many genera, particularly those of the Elina-series, 
are found in the temperate parts of South America. Three groups 
are recognized within the pronophilines: 

The Pronophila-series, sensu stricto, characterized by the gen- 
erally broad wings and rounded margins of the hindwings, except 
where modified by tails. The vast majority of these butterflies are 
found in tropical forests. 

The Elina-series, characterized by broad wings, but the hindwings 
are usually produced in the anal region. The forelegs are more 
reduced than in the Pronophi la-series. Most of these butterflies are 
found in the South Temperate region. 

The Lymanopoda-series, characterized by the narrow wings, 
rounded hindwing margins and reduced forelegs. Most of these 
species are found in the Andean highlands. 

The Pronophilini are characterized as follows: 

The eyes are usually hairy, but are naked in such diverse genera 
as Argyrophorus, Amphidecta, Auca, Spinantenna, Manerebia, Idio- 
neurula and Gyrocheilus. The antennae are generally two-fifths to 
half the length of the forewing costa. The antennal club is well 
developed, occupying the distal one-fifth to one-third of the antenna 
and between two and five times the thickness of the shaft. The third 
segment of the palpus is long for the Satyrini-section, never less than 
one-fourth as long as the second segment and frequently over half 
as long as the second in such genera as Lasiophila and Idioneurula. 



LEE D. MILLER 



11 




Figs. 261-272. Satyrinae: Pronophilini. 261. Pronophila thelebe Double- 
day and Hewitson. 6 venation. 262. P. thelebe, antennal club. 263. P. the- 
lebe, palpus. 264. Drucina leonata Butler, palpus. 265. Amphidecta pignera- 
tor Butler, palpus. 266. P. thelebe, 6 foreleg. 267. P. thelebe, 9 foretarsus. 
268. P. thelebe, midleg. 269. A. pignerator, midleg. 270. Pedaliodes poesia 
(Hewitson), 6 venation. 271. P. poesia, $ foreleg. 272. P. poesia, 2 fore- 
tarsus. 

The hairs on the second palpal segment are highly variable in their 
lengths: in such genera as Eteona and Dioriste the hairs are less 
than twice as long as the segment is wide, whereas in other genera, 



MEM. AMER. ENT. SOC, 24 



112 



THE SATYRIDAE 



such as Gyrocheilus and Manerebia, the hairs may be as much as five 
times the width of the segment. 

The male foreleg is moderately well developed in most genera 
with one (rarely two) tarsal subsegments that are devoid of spines 




Figs. 273-279. Satyrinae: Pronophilini. 273. Catargynnis pholoe (Stau- 
dinger), $ venation. 274. C. pholoe, palpus. 275. Coracles enyo Hewitson, 
8 venation. 276. Steroma bega Westwood, 6 venation. 277. Daedal ma 
dinias Hewitson, <$ venation. 278. D. dinias, palpus. 279. Argyrophorous 
argenteus Blanchard. c5 venation. 



LEE D. MILLER 



113 



(except in such genera as Cheimas, Gyrocheilus and Pronophila). 
The tibia is usually equal to, or longer than, the femur. Members 
of the Lymanopoda-series have greatly reduced forelegs with a mono- 
merous, unspined tarsus, and the femur is longer than the tibia. The 
female foreleg is usually moderately well developed with a penta- 
merus, clubbed tarsus bearing spines on the first four subsegments, 
but the foreleg of Gyrocheilus is greatly reduced with two unspined 
tarsal subsegments (Fig. 283). The relative lengths of the mid- 
and hindlegs are variable (Fig. 260). The mid-tibia is rarely less 
than twice as long as the first mid-tarsal subsegment, and it is gen- 
erally spiny dorsad, but smooth in such diverse genera as Amphidecta 
and Penrosada. The tibial spurs are present and well developed, 
and there is no spine at the dorsal, distal end of the midtibia. 

The forewing cell is usually excavate and half to three-fifths the 
length of the forewing costa. The forewing radial veins arise in 
three branches from the cell in most genera, but in such genera as 
Calisto, Steroma and Argyrophorus the radials arise in two, or even 
a single, stems; veins Rs and Mi arise connately (Idioneurula, etc.) 
to widely separate (as Gyrocheilus and Pedaliodes) . Vein M2 usually 





Figs. 280-283. Satyrinae: Pronophilini. 280. Proboscis propylea (Hew- 
itson), 6 venation. 281. Gyrocheilus patrobas (Hewitson), $ venation. 282. 
G. patrobas, palpus. 283. G. patrobas, 2 foretarsus. 



MEM. AMER. ENT. SOC, 24 



114 THE SATYRIDAE 

arises midway between Mi and Ms (but much nearer Mi in the 
Lymanopoda-series) , and Cui arises nearer M 3 than Cuo in some 
genera, midway between the two veins in some and nearer CU2 in 
others. Forewing vein Sc is always inflated, often greatly distended, 
and the other stalks may also be — all stalks are inflated in such 
genera as Steroma and Spinantenna, for example. 

The hindwing cell is produced by the distad displacement of 
m2-rri3 and half to over three-fifths the length of the wing measured to 
the end of M3. Vein Sc + Ri usually is longer than 3 A, and M3 and 
Cui are well separated at their origins. Vein M2 arises nearer, gen- 
erally much nearer, M3 than Mi. 

The pattern is variable and highly modified, and there is con- 
vergence between some of the Lymanopoda-series and the Holarctic 
Erebia. Most of the species are figured by Weymer, 1912 (1910- 
1912). 

Genera Included in the Pronophilini 2 

' Several genera have not been seen or were described too recently for 
critical inclusion in this revision. From the original descriptions it has, how- 
ever, been possible to place them in their taxonomic positions. Such genera 
are preceded by an asterisk (*). 

Pronophila-series, sensu stricto 

*Altopedaliodes Forster, 1964: 148. Type-species: Pronophila tena Hewitson, 

by original designation. 
Amphidecta Butler, 1867a: 404. Type-species: Amphidecta pignerator But- 
ler, by monotypy. 
*Antopedaliodes Forster, 1964: 151. Type-species: Pedaliodes antonia Stau- 

dinger, by original designation. 
Calisto Hiibner, 1816(1806-1838): 16. Type-species: Calisto zangis Hiibner, 

designated by Butler, 1868a: 194. 
Catargynnis Rober, 1892: 284. Type-species: Daedalma pholoe Staudinger, 

designated by Hemming, 1943: 23. 

=Pseudomaniola Rober, 1892: 222. Type-species: Daedalma pholoe 
Staudinger, by original designation. Preoccupied by Pseudomani- 
ola Weymer, 1890. 
Cheimas Thieme, 1906: 175. Type-species: Oxeoschistus opalinus Staudinger, 

by original designation. 
Corades Doubleday, 1848a: 115. Type-species: Corades enyo Hewitson, by 

monotypy. 
*Corderopedaliodes Forster, 1964: 155. Type-species: Pedaliodes corderoi 

Dognin, by original designation. 



LEE D. MILLER 



115 




Figs. 284-294. Satyrinae: Pronophilini. 284. Elina vanessoides Blanch- 
ard, $ venation. 285. E. vanessoides, palpus. 286. E. vanessoides, $ fore- 
leg. 287. E. vanessoides, 2 foretarsus. 288. E. vanessoides, midleg. 289. 
Spinantenna tristis (Guerin), 6 venation. 290. Lymanopoda samius West- 
wood, 6 venation. 291. L. samius, antennal club. 292. L. samius, palpus. 
293. Idioneurula erebioides (Felder and Felder), $ venation. 294. Manerebia 
cyclopina Staudinger, $ venation. 

MEM. AMER. ENT. SOC, 24 



116 THE SATYRIDAE 

Daedalma Hewitson, 1858(1856-1876): [85]. Type-species: Daeda'ma dinias 

Hewitson, designated by Butler, 18671: 268. 
Dioriste Thieme, 1906: 171. Type-species: Pronophila tauropolis Westwood, 

by original designation. 
Drucina Butler, 1872: 72. Type-species: Drucina leonata Butler, by monotypy. 
Eretris Thieme, 1905: 131. Type-species: Pronophila decorata Felder and 

Felder, designated by Hemming, 1943: 24. 
Eteona Westwood, 1850, in Doubleday, Westwood and Hewitson, 1846-1852: 

254. Type-species: Euterpe tisiphone Boisduval, by monotypy. 
Gyrochei'.us Butler, 1867f: 267. Type-species: Pronophila patrohas Hewitson, 

by original designation. This name was misspelled "Geirocheihts" by 

Holland, 1898: 211. 
Lasiophila Felder and Felder, 1859: 325. Type-species: Lasiophi'a cirta Felder 

and Felder, designated by Scudder, 1875a: 203. 
"Muscopedaliodes Forster, 1964: 153. Type-species: Pedaliodes muscosa 

Thieme, by original designation. 
Oxeoschistus Butler, 1867f: 268. Type-species: Pronophila puerta Hewitson, 

by original designation. 
Panarche Thieme, 1906: 228. Type-species: Corades tricordata Hewitson, by 

original designation. 
*Panyapedaliodes Forster, 1964: 157. Type-species: Pronophila panyasis 

Hewitson, by original designation. 
*ParapedaIiodes Forster, 1964: 153. Type-species: Pronophila parepa Hew- 
itson, by original designation. 
Pedaliodes Butler, 1867f: 267. Type-species: Pronophila poesia Hewitson, by 

original designation. 
*Pherepedaliodes Forster, 1964: 149. Type-species: Pedaliodes pheretiades 

Smith and Kirby, by original designation. 
*Physcopedaliodes Forster, 1964: 152. Type-species: Pronophila physcoa 

Hewitson, by original designation. 
Polymastus Thieme, 1906: 138. Type-species: Daedalma doraete Hewitson, 

by original designation. 
*Praepedaliodes Forster, 1964: 152. Type-species: Pronophila phanias Hew- 
itson, by original designation. 
*Praepronophi!a Forster, 1964: 182. Type-species: Pedaliodes emma Stau- 

dinger, by original designation. 
Proboscis Thieme, 1906: 168. Type-species: Pronophila propylea Hewitson, by 

original designation. 
Pronophila Doubleday and Hewitson, 1849, in Doubleday, Westwood and Hew- 
itson, 1846-1852: pi. 60. Type-species: Pronophila thelehe Doubleday 

and Hewitson, designated by Butler, 18671: 266. 

= Mygona Westwood, 1851, in Doubleday, Westwood and Hewitson. 
1846-1852: 357. Type-species: Pronophila thelehe Doubleday. 
designated by Hemming, 1939: 133. 
Pseudosteroma Weymer, 1912(1910-1912): 241. Type-species: Steroma pro- 



LEE D. MILLER 1 17 

nophila Felder and Felder, designated by Hemming, 1943: 25. 
*Punapedaliodes Forster, 1964: 148. Type-species: Pedaliodes albopunctata 

Weymer, by original designation. 
Steremnia Thieme, 1905: 137. Type-species: Pedaliodes (?) polyxo Godman 

and Salvin, designated by Hemming, 1943: 25. 
Steroma Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852: 

400. Type-species: Steroma bega Westwood, by monotypy. 
*Steromapedaliodes Forster, 1964: 148. Type-species: Pedaliodes albonolaia 

Godman, by original designation. 
Thiemeia Weymer, 1912(1910-1912): 267. Type-species: Pronophila phoro- 

nea Doubleday, designated by Hemming, 1943: 25. 



Elina-series 

Argyrophorus Blanchard, 1852: 30. Type-species: Argyrophorus argenteus 
Blanchard, by monotypy. 

Auca Hayward, 1953: 30. Type-species: Satyrus pales Philippi, by original 
designation. 

Elina Blanchard, 1852: 28. Type-species: Elina vanessoides Blanchard, desig- 
nated by Butler, 1868a: 194. 

*Haywardella Herrera, 1966: 71. Type-species: Satyrus thione Berg, by origi- 
nal designation. 

Nelia Hayward, 1953: 42. Type-species: Satyrus nemyroides Blanchard, by 
original designation. 

Neomaenas Wallengren, 1858: 78. Type-species: Neomaenas servilaea Wal- 
lengren, by monotypy. 

= Stibomorpha Butler, 1874: 204. Type-species: Stibomorpha deco- 
rata Butler (= Neomaenas servilaea Wallengren), by original des- 
ignation. 

Neosatyrus Wallengren, 1858: 79. Type-species: Neosatyrus ambiorix Wal- 
lengren, by monotypy. 

Pampasatyrus Hayward, 1953: 28. Type-species: Epinephele gyrtone Berg, 
by original designation. 

*Pseudomanio!a Weymer, 1890: 107. Type-species: Pseudomaniola euripides 
Weymer, designated by Hemming, 1943: 25. 

= Neomaniola Hayward, 1949: 156. Type-species: Pseudomaniola 
euripides Weymer, by original designation. Proposed to replace 
Pseudomaniola Weymer, wrongly believed to be preoccupied. 

*Quilaphoethosus Herrera, 1966: 69. Type-species: Satyrus monachus Blan- 
chard, by original designation. 

Spinantenna Hayward, 1953: 38. Type-species: Satyrus tristis Guerin, by 
original designation. 

Stuardosatyrus Herrera and Etcheverry, 1965: 74. Type-species: Argyro- 
phorus williamsianus Butler, by original designation. 

MEM. AMER. ENT. SOC, 24 



118 THE SATYRIDAE 

Lymanopo da-series 

Chillanella Herrera, 1966: 71. Type-species: Faunula stelligera Butler, by 

original designation. 
Cosmosatyrus Felder and Felder, 1867(1864-1867): 495. Type-species: Cos- 
mosatyrus leptoneuroid.es Felder and Felder, designated by Butler, 
1868b: 59. Herrera (1965: 70) lists the type of this genus as "Cos- 
mosatyrus chiliensis chiliensis C. et R. Felder", but chiliensis is a Guerin 
name. In any event, the type was properly designated by Butler. 
Etcheverrius Herrera, 1965: 62. Type-species: Satyrus chiliensis Guerin, by 

original designation. 
Faunula Felder and Felder, 1867(1864-1867): 488. Type-species: Faunula 

leucoglene Felder and Felder, by monotypy. 
Homoeonympha Felder and Felder, 1867(1864-1867): 487. Type-species: 

Homoeonympha pusilla Felder and Felder, by monotypy. 
Idioneurula Strand, 1942: 389. Type-species: Idioneura erebioides Felder 
and Felder, by original designation. 

= ldioneura Felder and Felder, 1867(1864-1867): 474. Type-species: 
Idioneura erebioides Felder and Felder, by monotypy. Preoccu- 
pied by Idioneura Selys, 1860. 
Lymanopoda Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 
1852: 401. Type-species: Lymanopoda samius Westwood, designated 
by Butler, 1868a: 196. 
Manerebia Staudinger, 1897: 139. Type-species: Manerebia cyclopina Stau- 

dinger, designated by Hemming, 1943: 24. 
Palmaris Herrera, 1965: 67. Type-species: Hipparchia monticolens Butler, by 

original designation. 
Penrosada Brown, 1944: 255. Type-species: Lymanopoda leaena Hewitson, 

by original designation. 
*Sabatoga Staudinger, 1897: 143. Type-species: Sabatoga mirabilis Stau- 
dinger, by monotypy. 
Sarromia Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 
1852: 402. Type-species: Sarromia obsoleta Westwood, by monotypy. 
Stygnolepis Strand, 1942: 389. Type-species: Sty gnus humilis Felder and 
Felder, by original designation. 

= Stygnus Felder and Felder, 1867(1864-1867): 487. Type-species: 
Stygnus humilis Felder and Felder, by monotypy. Preoccupied by 
Stygnus Perty, 1833. 
Tetraphlebia Felder and Felder, 1867 (1864-1867): 489. Typ;-species: Tetra- 

phlebia germainii Felder and Felder, by monotypy. 
Zabirnia Hewitson, 1877: 92. Type-species: Zabirnia zigomala Hewitson ( = 
Lymanopoda acraeida Butler), by monotypy. 

= Trophonina Rober, 1892: 222. Type-species: Lymanopoda acrae- 
ida Butler, by original designation. 



LEE D. MILLER 



119 



Tribe Satyrini Boisduval, 1836 

(Figures 295-308) 

Satyrides Boisduval. 1836: 166. 
= Hipparchiadae Kirby, 1837: 297. 

This tribe is at the center of its section, showing relationships 
with the Dirini by virtue of the general facies and the presence of 
the heavy spine at the dorsal, distal end of the midtibia, and with 
the Pronophilini and much of the Hypocystini in the shape of the 
hindwing cell. Through Davidina there is a progressive gradation 
from typical Satyrini to the Melanargiini. 



0.2xML, FL 



Satyrus 
Hipparchia 

Neominois 

Mmois 

Karanasa 

Davidina 

Oeneis 



Fig. 295. Satyrinae: Satyrini. Relative lengths of the femur + tibia + 
tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and 
hindlegs (HL, heavy stippling) of selected genera. In all instances the ML 
value is unity. The top bar for each genus represents the measurements ob- 
tained from males, the bottom bar those from females. 

The weakly developed forelegs' of both sexes serve to distinguish 
this tribe from the Hypocystini and much of the Pronophilini, and 
the production of the hindwing crossvein m-m a at M- will distinguish 
the satyrines from Melanargiini, Dirini and Erebiini. The most char- 




MEM. AMER. ENT. SOC, 24 



120 THE SATYRIDAE 

acteristic features, however, of the Satyrini are on the midleg: all 
genera except Chionobas and Oeneis have a heavy spine at the dorsal, 
distal end of the midtibia, and in all genera but the O^nm-series and 
Davidina the midtibia is less than one and one-fourth times as long 
as (and often shorter than) the first midtarsal subsegment. 

All members of the Satyrini are Holarctic. The Oeneis-series 
is found throughout the Holarctic, but the Satyrus-series is almost 
exclusively Palearctic: only Neominois is known from North America. 

There is a very interesting pattern similarity between the Euro- 
pean genus Minois and the North American Cercyonis, but while 
the European butterflies belong to the present tribe, Cercyonis is a 
member of the Maniolini. Several authors (e.g., Hemming, in press) 
have considered Cercyonis synonymous with Minois, but structural 
examination of the two genera does not confirm this conclusion, as 
noted in the discussion of Maniolini. 

Davidina is intriguing. Several authors have claimed it was a 
pierid, but examination of the tarsal claws shows Davidina to be a 
satyrid. Pierid tarsal claws are characteristically bifid (see Schatz 
and Rober. 1892), whereas those of Davidina are simple, as in most 
other satyrids. 

Two series are recognized in the Satyrini, as follows: 

The Satyrus-series, sensu stricto, distinguished by the extremely 
short midtibia and the presence of a heavy spine on the dorsal, distal 
end of the midtibia. 

The 0<?/7<?/s-series, distinguished by the longer midtibia lacking 
the spine at the dorsal, distal end. 

The Satyrini are characterized as follows: 

The eyes are naked. The antennae are more than two-fifths, but 
less than half, the length of the forewing costa. The antennal club 
is well developed, occupying the distal one-fourth to one-fifth of the 
antenna and three to more than five times as thick as the shaft (Fig. 
297). The third segment of the palpus is short, generally one-fourth 
to one-fifth as long as the second segment. The hairs of the second 
palpal segment are long, two and a half to five times as long as the 
segment is wide. 

The male foreleg is greatly reduced, the tibia and femur are of 
about the same length and the tarsus is mono- to trimerous without 
spines (Figs. 299, 306). The female foreleg is also greatly reduced, 
as much so as that of the male in several genera (Fig. 295), with 



LEE D. MILLER 



121 




301 




Figs. 296-303. Satyrinae: Satyrini. 296. Satyrus actaea (Linne), S 
venation. 297. S. actaea, antennal club. 298. S. actaea, palpus. 299. S. 
actaea, 6 foreleg. 300. S. actaea, 2 foretarsus. 301. S. actaea, midleg. 302. 
Brintesia circe (Fabricius), 6 venation. 303. Neominois ridingsii (Edwards), 
5 venation. 

the tarsus unspined, or only weakly spined, and comprised of one to 
four subsegments. The hindleg is significantly longer than the mid- 
leg (Fig. 295). In most genera the midtibia is less than one and 
one-fourth times the length of the proximal midtarsal subsegment, 
but in Davidina and the Oeneis-szrizs the midtibia is at least half 
again as long as the first midtarsal subsegment. The midtibia is very 
spiny dorsad, the spines being particularly heavy in this tribe, and 
the tibial spurs are present and well developed. There is a charac- 
teristic heavy spine at the dorsal, distal end of the midtibia in the 



MEM. AMER. ENT. SOC, 24 



122 



THE SATYRIDAE 



Satyrus-series (Fig. 301), which is absent in the Oeneis-series. 

The forewing cell is square-cut to slightly excavate and half to 
almost three-fifths the length of the forewing costa. The forewing 





Figs. 304-308. Satyrinae: Satyrini, all Oeneis noma (Thunberg). 304. 
$ venation. 305. palpus. 306. S foreleg. 307. 2 foretarsus. 308. midleg. 

radial veins arise from the cell in three branches, and veins Rs and 
Mi arise separate, but they may be approximate. Vein M 2 arises 
nearer Mi than M3, and Cui arises midway between M :) and CU2, 
or slightly nearer the latter. Forewing vein Sc is inflated basad, but 
the other veins are little, if at all, thickened. 

The hindwing cell is produced by the distad displacement of 
ni2-m 3 and half to three-fifths (slightly more in Oeneis) the length 
of the wing measured to the end of M3. Veins Sc + Ri and 3 A are 
of about the same length, and M :i and Cui are well separated at their 
origins. Vein M2 arises nearer Ms than Mi. 

The pattern is relatively uniform. The upper surface is shaded 
black, brown and/or orange and white with forewing ocelli in spaces 
M1-M3 and Cu 2 -2A. The under surface pattern repeats the forewing 
ocelli, and the hindwings are cryptically marked. Davidina is white 
or off-white with dark brown to black veinal and interveinal lines. 
Illustrations of many of the species are given by Moore, 1893(1890- 
1893). 



LEE D. MILLER 123 

Genera Included in the Satyrini 
Satyrus-series, sensu stricto 

Arethusana de Lesse, 1951: 40. Type-species: Papilio arethusa Esper, by 

original designation. 
Aulocera Butler, 1867b: 121. Type-species: Satyrus brahminus Blanchard, 

designated by Butler, 1868a: 194. 
Berberia de Lesse, 1951: 41. Type-species: Satyrus abdelkader Pierret, by 

original designation. 
Brintesia Fruhstorfer, 1912(1912-1915): 307. Type-species: Papilio circe 

Fabricius, by original designation. 
Chazara Moore, 1893(1890-1893): 21. Type-species: Papilio briseis Linne, 

by original designation. 
Davidina Oberthiir, 1879: 19. Type-species: Davidina armandi Oberthur, by 

monotypy. 
Eumenis Hiibner, [1819]( 1816-1826) : 58. Type-species: Papilio autonoe 

Esper, designated by Grote, 1873: 62. 
Hipparchia Fabricius, 1807: 281. Type-species: Papilio fagi Scopoli, desig- 
nated by Butler, 1868a: 194. 

=Nytha Billberg, 1820: 77. Type-species: Papilio hermione Linne 

(=Papilio fagi Scopoli), designated by Scudder, 1875a: 231. 
=Melcmia Sodoffsky, 1837: 81. Type-species: Papilio fagi Scopoli, 
by original designation. Preoccupied by Melania Lamarck, 1799. 
Proposed to replace Hipparchia Fabricius, wrongly believed to be 
preoccupied. 
Kanetisa Moore, 1893(1890-1893): 42. Type-species: Hipparchia digna 

Marshall, by original designation. 
Karanasa Moore, 1893(1890-1893): 38. Type-species: Satyrus huebneri Feld- 

er, by original designation. 
Minois Hiibner, [18 19] ( 1816-1826) : 57. Type-species: Papilio phaedra 

Linne (=Papilio dryas Scopoli), designated by Butler, 1868a: 194. 
Neohipparchia de Lesse, 1951: 40. Type-species: Papilio statilinus Hufnagel, 

by original designation. 
Neominois Scudder, 1875b: 241. Type-species: Satyrus ridingsii Edwards, by 

original designation. 
Paroeneis Moore, 1893(1890-1893): 36. Type-species: Chionobas pumilus 

Felder, by original designation. 
Philareta Moore, 1893(1890-1893): 23. Type-species: Papilio hanifa Nord- 

mann (=Papilio anthe Ochsenheimer), by original designation. 
Pseudochazara de Lesse, 1951: 42. Type-species: Satyrus pelopea Klug, by 

original designation. 
Pseudotergumia Agenjo, 1947: unnumbered page at end. Type-species: Pa- 
pilio phidia Linne, by original designation. 
Satyrus Latreille, 1810: 355. Type-species: Papilio actaea Linne, designated 

MEM. AMER. ENT. SOC, 24 



124 



THE SATYRIDAE 



by the action of the International Commission on Zoological Nomen- 
clature, 1943. Opinion 142. 

Oeneis-series 

Chionobas Boisduval, [1833]( 1832-1843) : 182. Type-species: Papilio aello 
Hiibner (=Papi!io glacialis Moll), designated by Blanchard, 1840: 457. 

Oeneis Hiibner, [ 1819] ( 1816-1 826) : 58. Type-species: Papilio noma Thun- 
berg, designated by Butler, 1868a- 196. 

Tribe Melanargiini Verity, 1920 
(Figures 309-314) 

Melanargiinae Verity, 1920: 56. 
= Agapetinae Verity, 1953: 3, 46. 

This tribe contains some of the most distinctive of all satyrids, 
the pattern being unmistakable. In the shape of the hindwing cell, 
morphology of the legs and general facies (excluding the white ground 



0.2 x ML, FL 




Melanargia 
Ledargia 

Argef ormia 



Fig. 309. Satyrinae: Melanargiini. Relative lengths of the femur + 
tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip- 
pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances 
the ML value is unity. The top bar for each genus represents the measure- 
ments obtained from males, the bottom bar those from females. 

color), these butterflies closely resemble the Dirini and probably 
developed from a common stock with them. The spiny legs, reduc- 
tion of the forelegs and other structures suggest a strong affinity of 
the melanargiines and the Satyrini. There is little to relate this tribe 
closely either to the Hypocystini or to the pronophilines. 

The pattern of these butterflies sets them apart from all relatives 
— white with black marbling and well developed sets of ocelli on 
both wings, particularly the hindwings. 

All members of the Melanargiini are found in the Palearctic from 



LEE D. MILLER 



.25 



Europe to China and Manchuria. There are more species in Asia 
than in Europe. The Melanargiini are characterized as follows: 

The eyes are naked. The antennae vary little from half the 
length of the forewing costa. The antennal club is rather weakly 
developed, less than two to less than three times the thickness of 
the shaft. The third segment of the palpus is one-third to one-fourth 
the length of the second segment. The hairs of the second segment 
of the palpus are about four times as long as the segment is wide. 

The male foreleg is greatly reduced, with a single, unspined tarsal 
subsegment; the femur is longer than the tibia (Fig. 312). The 
female foreleg is as reduced as that of the male with a monomerous, 
unspined tarsus (Fig. 313). The relative reduction of the fore-, mid- 
and hindlegs is shown in Fig. 309. The midlegs are usually shorter 
than the hindlegs (Fig. 309). The midtibia ranges from just under 
to just over one and a half times as long as the proximal midtarsal 






Figs. 310-314. Satyrinae: Melanargiini, all Melanargia galathea (Linne). 
310. 6 venation. 311. palpus. 312. 6 foreleg. 313. 2 foretarsus. 314. 
midleg. 

subsegment and is very spiny dorsad. The tibial spurs are present 
and well developed; there is no heavy spine at the dorsal, distal end 
of the midtibia. 

The forewing cell is square-cut and approximately half the length 
of the forewing costa. The forewing radial veins arise in three 
branches from the cell, and veins Rs and Mi are separate, but may 



MEM. AMER. ENT. SOC, 24 



126 THE SATYRIDAE 

be approximate, at their origins. Vein M 2 arises much nearer Mi 
than M3, and Cui arises nearer M3 than CU2. If any forewing veins 
are inflated, only Sc would be, and that not extensively. 

The hindwing cell is produced at the origin of M3 and half to 
three-fifths the length of the wing measured to the end of Ms. Veins 
Sc + Ri and 3 A are of about the same length, and M3 and Cui are 
well separated at their origins. Vein M 2 arises slightly nearer Mi 
than M3. 

The pattern is quite distinctive: all species are white or cream- 
colored marbled with black and with the ocelli well developed. Many 
genera have been named, but it seems that one genus would suffice 
biologically for all the species. 

Genera Included in the Melanargiini 

Argeformia Verity, 1953: 47. Type-species: Papilio arge Sulzer, by original 
designation. Proposed as a subgenus of Agapetes Billberg (=Mela- 
nargia Meigen). 
Epimede Houlbert, 1922: 132, 142, 160. Type-species: Arge halimede Mene- 

tries, designated by Hemming, 1934a: 143. 
Halimede Oberthur and Houlbert, 1922: 192. Type-species: Halimede asiatica 

Oberthiir and Houlbert, by monotypy. 
Lachesis Oberthur and Houlbert, 1922: 192. Type-species: Lachesis rusci- 
nonensis Oberthur and Houlbert ( =Arge lachesis Hiibner), by original 
designation. 
Ledargia Houlbert, 1922: 157. Type-species: Melanargia leda Leech, by 

original designation. 
Melanargia Meigen, 1829: 97. Type-species: Papilio galathea Linne, desig- 
nated by Kirby, 1894: 41. 

=Arge Hiibner, [1819]( 1816-1826) : 60. Type-species: Papilio 
psyche Hiibner (= Papilio occitanica Esper), designated by Butler, 
1868a: 196. Preoccupied by Arge Schrank, 1802. This name 
was suppressed in favor of Melanargia Meigen by the International 
Commission on Zoological Nomenclature, 1956, Opinion 400. 
= Agapetes Billberg, 1820: 78. Type-species: Papilio galathea Linne, 
designated by Scudder, 1875a: 104. This name was suppressed in 
favor of Melanargia Meigen by the International Commission on 
Zoological Nomenclature, 1956, Opinion 400. 
Parce Oberthur and Houlbert, 1922: 193. Type-species: Parce fergana Ober- 
thiir and Houlbert (=Melanargia parce Staudinger), by original des- 
ignation. 



LEE D. MILLER 



127 



Genera of Uncertain Position 

Genus Pamperis Heimlich, 1959 



(Fig. 315) 



Pamperis Heimlich, 1959: 177. Type-species: Pamperis poaoeneis Heimlich, 
by original designation. 




Fig. 315. Venation of Pamperis poaoeneis Heimlich (after Heimlich, 
1959). Note the aberrant venation pattern discussed in the text. 

Having seen no specimens of the type-species of this Neotropical 
monotypic genus, I can only evaluate its position in terms of the 
original description, which leaves many questions unanswered. Heim- 
lich's photographs of the types suggest he was correct, at least 
superficially, in placing this genus near Cosmosatyrus Felder and 
Felder, and as such, it certainly belongs in the Lymanopoda-seriQs 
of the satyrine tribe Pronophilini. However, the venational drawing 
given represents either an extraordinary aberration — the veins in the 
apical portion of the forewing are arranged like no known satyrid, 
and the hindwing lacks a major vein (Fig. 315) — or represent draft- 
MEM. AMER. ENT. SOC, 24 



128 THE SATYRIDAE 

ing errors. Until these structures, as well as others not considered 
in the original description, have been studied, it will be quite im- 
possible to assign this genus to its proper position among the Satyridae. 

Genus Setodocis Billberg, 1820 

Setodocis Billberg, 1820: 78. Type-species: Papiiio periboea Fabricius, de- 
signated by Scudder, 1875a: 268. 

I have no idea to what this genus refers, and neither does anyone 
else; periboea is a "lost" species. The best opinion is that of Hem- 
ming (in press), "Include at the end of the subfamily Mycalesinae as 
a genus dubium." The name is in the literature, so it must be men- 
tioned here, but it would seem advisable to suppress it. 

The Evolution and Zoogeography of the Satyridae 

The twin subjects of evolution and biogeography have provided 
biologists since the days of Charles Darwin and Alfred Russel Wal- 
lace with the raw material for sundry mental gymnastics and manipu- 
lations of the earth's flora, fauna, history and even crust through 
space and time. Such exercises, while based on the best available 
evidence, are limited by that evidence, and what are proposed as facts 
must be considered in actuality only as more or less educated guesses. 
The preceding statement is particularly true when one is dealing with 
such insects as butterflies, the fossil records of which are almost non- 
existent. In drawing evolutionary, and especially zoogeographic, in- 
ferences it is necessary to guess the past history of butterflies by com- 
parison with fossil histories of vertebrates and to interpret tendencies, 
such as the simplification of structures through phylogenetic lines, by 
the interpretations given such trends in groups with reasonably docu- 
mented fossil records. 

Zoogeography, in particular, has been a constant source of 
controversy, with several opposing schools of thought attempting to 
analyze distributional data in accordance with conflicting theories. 
At present there are two major camps of biogeographers which may be 
called the Wegener and the Matthew schools, although each theory 
has been refined far beyond its originator's proposals. 

"Nothing in zoogeography has brought forth more argument or 
more demands for an open mind — the other man's mind — than the 
idea of continental drift." (Darlington, 1957: 606). This theory, 



LEE D. MILLER 129 

first put forth by Wegener, states briefly that the continents were a 
single landmass which broke apart, and the pieces drifted across the 
relatively plastic mantle, chiefly during the Mesozoic and into the 
Tertiary, to their present positions. By rotating the present-day 
continents and fitting them together it is possible to arrive at a con- 
vincingly completed "jig-saw puzzle"; indeed, it was this sort of 
piecing together of the landmasses of the world which provided the 
basis for the theory in the first place. Caster (1952) summarized 
the stratigraphic evidence supporting connections between South 
America and Africa, particularly during the Mesozoic, and further 
cited the similarity between the shallow water invertebrates of the 
two continents, but Dunbar ( 1952: 154) demonstrated by the simi- 
larity of shallow water molluscs of Kwajalein, Bikini and Guam that 
many shallow water invertebrates can cross deep water "barriers", 
probably as free-swimming larvae. Darlington (1957: 607-608) 
showed another of the problems with the acceptance of the Wegener 
hypothesis as an explanation of the present-day animal distributions, 
when he stated, ". . . if a group of animals now occurs only in 
Africa and Brazil, it is said to date from and be evidence of a hypo- 
thetical African-Brazilian continent. This method depends on an 
assumption which Wegenerians usually do not put into words: that 
animal distributions are more permanent than land, that animals 
move less than continents." This argument more or less summarizes 
my own objections to continental drift as an explanation for the dis- 
tributions of present-day organisms. 

Most American zoogeographers today subscribe to the general 
theory of animal dispersal proposed by Matthew (1915) and elab- 
orated upon by Hesse, Allee and Schmidt (1951) and Darlington 
(1957). Matthew set forth his thesis in the beginning of his paper 
(1915: 173). 

While the details of Matthew's work have been refined, the 
general skeleton has remained intact. Geologic evidence, such as that 
presented by Schuchert (1935) for the history of the Neotropics, 
has strengthened the Matthew theory. Biogeographers have been 
better able to devote their attention to the more knotty problems of 
how, when and why organisms arrived where they are. 

Since Schuchert (1935) showed the existence of several Ter- 
tiary "water gaps" between North and South America and roughly 

MEM. AMER. ENT. SOC, 24 



130 THE SATYRIDAE 

dated them, many authors have considered it an absolute necessity 
to close these gaps physically so that organisms might cross them. 
This practice, however, has recently come under scrutiny from sev- 
eral authors, notably Simpson (1952), who demonstrated mathe- 
matically that given a length of time of sufficient duration and a 
sufficiently large "feeder" population even the most improbable dis- 
persal of a terrestrial organism over water could be a virtual certainty. 
While this mathematical exposition does not serve as evidence of a 
population of an area via waif dispersal, it shows a method by which 
such dispersal could be accomplished. It is of great importance to 
the understanding of the aerial dispersal of such insects as butterflies 
because it shows that landmasses need never be physically connected 
to exchange faunal elements across barriers. Such studies indicate 
that reappraisals are in order concerning the necessity of erecting 
land bridges across ocean deeps. Certainly physical barriers work 
less hardship on such strong flying butterfly genera as Danaus (Da- 
naidae), Precis and Vanessa (Nymphalidae), Libythea (Libytheidae) 
and Leptotes and Lampides (Lycaenidae) than on the more sedentary 
Satyridae and Ithomiidae. 

I wish here to call attention to the second paragraph of Matthew's 
thesis (1915: 173). Nearly all the principal routes of migration pass 
through either the eastern Palearctic or the Indo-Malayan region. 
If a group is in a rather generalized condition at the time of its 
expansion, it seems illogical that such a group would not leave some 
more or less primitive members along its primary path of dispersal. 
In the case of biotic interchange between the Palearctic and the 
Nearctic, the Paleotropics and the Neotropics, Asia and Australia, 
Africa and Australia, these waves of dispersal would have extended 
through eastern Asia. Eastern Asia also provides the complete spec- 
trum of biotopes, from Arctic tundra to tropical rainforest to the 
most arid desert — a prospective "home" for almost any group. No 
other region on earth is fed by so many major routes of dispersal, 
hence, no other region should be expected to have organisms derived 
from so many different sources. This diversity has led many students 
(e.g., Darlington, 1957) to consider eastern Asia the major center 
of dispersal, a veritable "Garden of Eden". In the following pages 
I shall attempt to show that at least the Satyridae need not have 
arisen in this region, although eastern Asia certainly played an 



LEE D. MILLER 131 

important role in the subsequent evolution of the family. 

Origin of the Satyridae 

Consideration of the origin and subsequent evolution of the 
Satyridae must be prefaced by a discussion of their relatives. Two 
nymphaloid families, the Morphidae (including the Amathusiidae) 
and Ithomiidae, are closely allied to the family under consideration 
here and have been characterized at the beginning of the systematic 
revision. The evolution of all three families must be considered 
together to lay the groundwork for that which follows. 

The living nymphaloid families divide basically into two major 
groups on larval foodplant requirements. With very few exceptions 
(Ehrlich and Raven, 1965) the "true" nymphalids feed on dicoty- 
ledonous plants, whereas the larvae of the satyrids utilize monocoty- 
ledons exclusively. 

Forbes (1932) has assumed that the modern lepidopterous fami- 
lies had already differentiated by the Jurassic, and Carpenter (1930) 
felt that the Lepidoptera as an order could be traced only as far 
back as the Jurassic. Present evidence (Just, 1948) indicates that 
the proliferation and differentiation of the angiosperms took place 
in the late Mesozoic. The differentiation of the nymphaloid stock 
must have been somewhat synchronous with this proliferation of the 
flowering plants, but there is some question about this point. Fox (in 
lift.) feels that there is no reason to assume that morphological 
differentiation may not have preceded plant evolution, i.e., present 
monocotyledon feeders may have begun as dicotyledon feeders, or 
vice versa. Whether plant or butterfly evolution came first becomes 
a moot point, but the potential new niche provided by the evolution 
of the two basic stocks of plants must have provided some impetus 
for the subsequent, if not the initial, evolution of the butterflies. In 
any event, the butterflies are a relatively recent group compared with 
the Orthoptera and the Odonata, which were well developed by the 
Carboniferous. 

The Ithomiidae feed as larvae almost exclusively on the So- 
lanaceae (Fox, 1956: 13) and belong on this basis to the "true" 
nymphalids, although they are not far removed from the satyrid 
complex. These butterflies have reached their highest development 

MEM. AMER. ENT. SOC, 24 



132 THE SATYRIDAE 

in the Neotropics, although a single tribe, the Tellervini, is found in 
Australia and New Guinea. 

The Satyridae, sensu stricto, overwhelmingly feed as larvae on 
Gramineae and Cyperaceae, and all known satyrids feed on mono- 
cotyledons. Some members of the family feed on Palmaceae (Elym- 
nias and relatives). All known brassolines feed on monocots, 
generally utilizing the primitive ones, but a few also feed on bamboo; 
none are known to use dicots. The brassolines are exclusively Neo- 
tropical, none living farther north than Mexico. The foodplant 
requirements of the other primitive Neotropical satyrids (Haeterinae 
and Biinae) are unknown, but since they are restricted to the deep 
forest they probably do not use grasses, other than possibly bamboo. 
There are many other monocotyledons in the rainforest, and these 
are probably the food plants. 

Structurally, the Indo-Australian Amathusia, Faunis, Taenaris, 
etc. belong with the American Morpho in the family Morphidae. All 
the known Indo-Australian Morphidae feed on monocotyledons as 
larvae, many using Gramineae. In the New World, however, the 
morphids (represented only by the nominate genus) predominantly 
feed on dicotyledonous plants of the family Canellaceae, Myrtaceae, 
Menispermiaceae, Rhamnaceae, Sapindaceae and Leguminoseae. 
One species, however, Morpho aega (Fabricius), feeds on bamboo 
(Gramineae). Morpho is one of the few nymphaloid genera known 
which utilizes both monocotyledons and dicotyledons as larval food; 
indeed, the Morphidae is the only family commonly showing such 
habits. It seems evolutionary significant that the genus Morpho is 
limited to the New World tropics. Furthermore, excluding the 
brightly colored and highly modified upper surface of these butter- 
flies, the morphids seem to typify the "primitive nymphalid pattern" 
of Schwanwitsch (1924): most of the elements of the pattern are 
present in the primitive species. 

In summary, many of the most primitive members of various 
nymphaloid families occur only in the Neotropics. The Neotropical 
tribe Tithoreini of the Ithomiidae is as primitive as the Indo- 
Australian Tellervini, and the two might well be combined were it 
not for their geographic isolation and a few morphologic characters 
(Fox, 1956: 20). As shown above, the Neotropical genus Morpho 
is the most primitive of the morphids. The most primitive satyrids 
are the Neotropical Haeterinae, which preserve the forewing vein 3A. 



LEE D. MILLER 



133 




Fig. 316. The early distributional history of the Haeterinae, Brassolinae 
and Biinae in the New World. All movement illustrated in this figure was 
during the Cretaceous and earliest Tertiary (open arrows). In succeeding 
figures stippled arrows indicate mid-Tertiary movement, whereas solid black 
arrows indicate late Tertiary and Quaternary migration. Major areas of the 
world are indicated on this map for reference, and no attempt is made to 
present exact continental outlines at any given period of geologic time. 

The next most primitive groups of satyrids are the biines and the 
brassolines and the majority of these, including the most primitive 
ones, are New World species. In addition to these satyrids, the 
most primitive danaids (Clothilda and the Lycoreinae) are found in 
Central America and the Antilles. The remaining danaids are de- 
rived from Paleotropical groups which arrived in the Neotropics 
much later. 

In view of the evidence compiled in the preceding paragraphs, a 
feasible explanation of the evolution of the nymphaloid butterflies 
seems to require serious consideration of tropical America as some- 
thing other than a receptacle for pre-existing Holarctic groups. Prob- 
ably a relatively undifferentiated nymphaloid stock either entered the 
Neotropics as part of the extensive Holarctic invasion during the 

MEM. AMER. ENT. SOC, 24 



134 THE SATYRIDAE 

Cretaceous (Darlington, 1957: 561-564), or this stock was already 
in tropical America prior to the connection of South America with 
the Holarctic. Since there are so many highly primitive groups of 
the Satyridae, Morphidae, Ithomiidae, and Danaidae in the Neo- 
tropics, it further appears that it was in tropical America that the 
fundamental bifurcation of the nymphaloid stock occurred. At least 
these four families appear to have arisen in the Neotropics. Later 
all four reinvaded the Holarctic, probably during the Cretaceous, but 
only two of them (Satyridae and Danaidae) reinvaded the American 
tropics during the Tertiary. The Tertiary history of the danaids 
probably parallels that of the satyrids, which will be discussed later. 
Endemic, primitive, though often highly specialized, stocks of all 
four families may be still found in the Neotropics. It seems most 
likely, therefore, that these families all arose in the Cretaceous (pos- 
sibly earlier) rainforests of tropical America. 

The Later History of the Satyridae 

The most primitive satyrids, the Haeterinae, are now restricted 
to the Neotropical rainforest, and there is no indication that they 
ever were elsewhere, and all evolution in this group has occurred 
within the American tropical forests (Fig. 316). No members en- 
tered the Old World, as did members of the Ithomiidae. It is also 
doubtful that these insects ever exploited the grasslands that resulted 
from progressive drying during the Tertiary. In fact, these insects 
are singularly well adapted to the deep forest — the hyaline wings of 
many species and the ghost-like fluttering flight render them virtually 
invisible except when they pass through patches of sunlight. Masters 
(in lift.) says that Haetera always restricts itself to the deepest forest, 
never entering sunlit areas, but I have seen species of Pierella and 
Callitaera sunning themselves early in the morning, but not later 
in the day. 

Figs. 317, 318. Fig. 317. The origin and subsequent distributional his- 
tory of the Melanitini (Biinae) and the origin of the Elymniinae from the 
Melanitini. For details, see text. Fig. 318. The origin and proliferation of 
the Elymniinae into the tribes Lethini (see Fig. 319), Elymniini (Fig. 320), 
Mycalesini (Fig. 321) and Zetherini. The latter tribe did not significantly 
leave the Indo-Malayan region. The probable origin of the Eritinae is also 
shown on this map. For details, see text. 



LEE D. MILLER 



135 




ftethini (Fig. 319) 

Elymniini(Fig. 320) 
Elymniinae / 

1 \\ Mycalesini (Fig. 321) 

>•< V 7etherini 





MEM. AMER. ENT. SOC. 24 



136 THE SATYRIDAE 

The history of the Brassolinae probably closely parallels that of 
the Haeterinae (Fig. 316). These butterflies, however, are strong 
fliers and have evolved a "flash coloration pattern" on the upper 
surface and a cryptic under surface. They do not depend so much 
on camouflage as on the strength of their flight. Many species are 
crepuscular, particularly those occurring in more open country, but 
most species are more or less restricted to the rainforest. 

The Biinae did move into the Holarctic and apparently provided 
the basis for the evolution of the rest of the family. Those biines 
which did not leave the Neotropics (Fig. 316) have remained rain- 
forest butterflies with patterns and flight characteristics similar to 
those of the brassolines. 

Whether the Neotropical members of any of the preceding sub- 
families ever invaded the grasslands is questionable. If they did, 
they were wiped out, probably in competition with the better adapted 
Satyrinae which later came into the New World. It seems more 
likely, however, that these butterflies never left the rainforest of 
their origin and evolved into such ecologically successful species that 
they easily resisted competition from the satyrines; perhaps these pri- 
mitive groups actually forced the satyrines into more open country. 

Those Biinae which did move into the Holarctic were obviously 
much more like Antirrhini than Bia, but there are now no New World 
antirrhines that have extremely close relationships with the Old World 
species. The migrating biines were forced into the Paleotropics at a 
fairly early time, where they apparently evolved into the present- 
day tribe Melanitini ( Fig. 317). Several of the melanitines have 
become very widespread, apparently relatively recently, and one, 
Melanitis leda (Linne), is now found from West Africa to Mada- 
gascar and east into Australia and Japan. Most of the species are 
found in the Indo-Malayan region and in tropical Africa (Table 7). 
The Melanitini responded to the increase in aridity of the early 
Tertiary by invading open woodlands and even more open country, 
being thus the first satyrids, historically and evolutionarily, to leave 
the deep forest. These butterflies are more or less crepuscular, like 
the Brassolinae. A possible explanation for the crepuscular flight 

Figs. 319, 320. Fig. 319. The distributional history of the Lethini. See 
text for details. Fig. 320. The distributional history of the Elymniini. See 
text for details. 



LEE D. MILLER 



137 




MEM. AMER. ENT. SOC, 24 



138 



THE SATYRIDAE 




Fig. 
details. 



321. The distributional history of the Mycalesini. See text for 



habit is that such species are actually approximating the conditions 
of the deep forest as regards light intensity. In short, such species 
are basically forest species that are "carrying the forest environment 
with them". The successful invasion of the open country provided 
the impetus for the great adaptive radiation and phylogenetic diversi- 
fication of the satyrids in the Old World. The Neotropical Antirrhini, 
on the other hand, never became successfully adapted to open country, 
and without leaving the forests evolved into the present tribe with 
fewer than twenty species. In the Indo-Australian region the Melani- 
tini were probably giving rise, meanwhile, to the Elymniinae through 
steps resembling the present-day genera Parantirrhoea (Melanitini), 
Pty chandra and Samanta (both Lethini), shown in Figs. 317, 318. 

The Elymniinae proliferated rapidly during late Cretaceous and 
early Tertiary into the four tribes, most of which stayed in the wood- 
lands and scrublands: some (a number of Mycalesini) invaded the 
savannas, and others (the Neorina-series) re-entered the deep forests. 
The pattern of early and later migration of the subfamily rather closely 
parallels that of many vertebrate groups and seems to have been 



LEE D. MILLER 139 

limited northward by the Tethys Sea. Little movement from the 
Paleotropics seems to have been accomplished before the closing of 
this "water gap" between the Paleotropics and the Palearctic (Fig. 
318). Interrelationships within the Elymniinae can be better drawn 
than those within any other subfamily. The Lethini grade toward the 
Zetherini, on the one hand, and the Mycalesini (through such genera 
as Mandarinia and Orsotriaena), on the other, as brought out in 
the systematic revision. The Mycalesini connect well with the Elym- 
niini, as do the zetherines. The elymniines are "throwbacks'"' in one 
respect: all known members feed as larvae on Palmaceae, hence the 
common name "palmflies". All tribes of the Elymniinae but the 
Zetherini are widely distributed in the Old World at present, although 
only one — the Pararge-section of the Lethini — has managed signifi- 
cantly to penetrate the Palearctic. The distributional histories of the 
tribes are shown in Figs. 319 (Lethini), 320 (Elymniini), 321 
(Mycalesini); the actual patterns are discussed in the analyses of the 
individual faunae of the world. 

Apparently within a short time — probably still in the Cretaceous 
— the Elymniinae gave rise to the Satyrinae (Figs. 318, 322), pre- 
sumably through either the Mycalesini or (more probably) the 
Lethini and the Hypocystini of the Satyrinae. It is also probable that 
the Eritinae arose from a basic elymniine stock (Fig. 318) and pos- 
sible that the Ragadiinae did (Fig. 322), but the origin of the latter 
subfamily is among the most obscure in the family Satyridae. 

The hypocystine genus Lamprolenis, while a fairly typical member 
of that tribe, retains some very primitive characteristics similar to 
those of the Mycalesini (Fig. 148). There is a typical Mycalesis- 
like hair tuft on the hindwing, and the general facies embody a com- 
bination of mycalesine and hypocystine characteristics. 

The other case of similarity between the Elymniinae and Satyrinae 
is to be found in South Africa, where members of the Aeropetes-series 
of the Lethini seem to grade into the Dirini (Figs. 67-72 and 248-259, 
respectively). It would be tempting to ascribe great evolutionary 
significance to the similarity, but the A eropetes-series contains primi- 
tive lethines, whereas the Dirini appear to be fairly advanced Sa- 
tyrinae — particularly as regards the reduction of the forelegs of both 
sexes. Therefore, this similarity is ascribed to convergence, not to 
phyletic proximity. 

MEM. AMER. ENT. SOC, 24 



140 THE SATYRIDAE 

The strongest link between the Elymniinae and the satyrines 
occurs through the subfamily Eritinae. Eritines have many charac- 
teristics in common with the Lethini, as discussed in the systematic 
revision, and also share characters with the Hypocystini. This can- 
not be merely convergence, and it appears that the Satyrinae arose 
in the Indo-Australian region from an elymniine stock not far re- 
moved from the lethine stock (Figs. 318, 322). 

In any event, the Satyrinae — probably from a Hypocystini-like 
stock — soon split into two basic groups, the Ypthimini- and Satyrini- 
sections (Fig. 322). This dichotomy was probably brought about 
by the increasing aridity and proliferation of savannas during the 
early Tertiary. The Ypthimini-section generally occupies more open 
country than do the Satyrini-section butterflies. 

During the early to mid-Tertiary the various tribes of the Satyri- 
nae, except the Pronophilini and Euptychiini, probably evolved in 
the Old World (Figs. 323-326). Upon the reopening of the Cen- 
tral American land connection between North and South America — 
it was probably sufficiently opened in Miocene, perhaps Oligocene, 
times — the forerunners of the principal South American tribes of 
the Satyrinae, the Pronophilini (derived from the Satyrini-section, 
Fig. 325) and the Euptychiini (from the Ypthimini-section, Fig. 
323 ) reentered South America and rapidly assumed dominance over 
the "old" South American fauna. At least part of this dominance 
may be ascribed to the ecological diversity shown by both tribes, as 
explained in the discussions of the tribes in the systematic revision. 
A glance at the numbers of species involved in both the Pronophilini 
and Euptychiini shows the dramatic radiation that has taken place in 
the Neotropics. 

It is doubtful that the radiation of the Euptychiini and Pronophi- 
lini resulted in a wholesale extinction of the Haeterinae, Brassolinae 
and Biinae, since the new arrivals probably did not compete ecologi- 
cally with the already established primitive ones. Both the Pro- 
Figs. 322, 323. Fig. 322. The origin of the Satyrinae from the Elymniinae 
and the division of the satyrines into the Hypocystini, the Ypthimini-section and 
the Satyrini-section. The probable origin of the Ragadiinae from the Elym- 
niinae is also shown in this figure. For details, see text. Fig. 323. The origin 
and distributional history of the Ypthimini (Old World) and Euptychiini (New 
World). See text for details. 



LEE D. MILLER 



141 



Ypthimin i-sect_ 

Satyrini-sect.f Satyrinae 

o 

elymniine ancestor 





Ragadiinae 



Hypocystini 



MEM. AMER. ENT. SOC, 24 



142 



THE SATYRIDAE 




Fig. 324. The origin and distributional history of the Coenonymphini, 
Maniolini and Erebiini. See text for details. 

nophilini and Euptychiini feed as larvae predominantly on Grami- 
neae, which are probably not used extensively by the primitive species. 

Just before and during the Pleistocene there was an exchange 
of faunal elements between the Nearctic and the Palearctic, resulting 
in such Holarctic distributions as those of Coenonympha (Fig. 324), 
Oeneis (Fig. 326) and Erebia (Fig. 324), although these genera 
probably differentiated much earlier, and some of the Holarctic mem- 
bers of the Maniolini (Fig. 324) and Satyrini (Fig. 326). 

A proposed phylogeny for the Satyridae is presented in Fig. 327, 
embodying the information given above. 



Derivation of the Satyrid Faunae of the World 

Comparisons of the faunal compositions of various parts of the 
world often prove valuable in determining the derivation of the 
elements of some areas. With this assumption in mind Table 7 was 
drawn to show the composition by tribes of the Satyridae in major 
faunal regions of the world. 



LEE D. MILLER 



143 




Figs. 325, 326. Fig. 325. The origin and distributional history of the 
Dirini and Pronophilini. See text for details. Fig. 326. The origin and dis- 
tributional history of the Satyrini and Melanargiini. See text for details. 



MEM. AMER. ENT. SOC, 24 



144 



THE SATYR1DAE 



Cretaceous 



TIME approx. 



early Tertiary 



TAXON 



mid-Tertiary 



Tribe 



Subfamily 



Melanargiini _ 

Satyrini 

Ptonophilini 

Ditlni 

Hypocystini 

Erebiini 

Maniolinl 

Coenonymphini 

Ypthimini 

Euptychiini '' 



>SATYRINAE 



Eritlni 

Mycalesin J-___ 
Ely m niini 
Zetherini 
Lethini- — 

Melanitini 

Antirrhini 

Biini 

Bfassolini 

Haeterini 



-RAGADIINAE 
— ERITINAE 



_~~~-ELYMNIINAE 



_.;::: BIINAE 

-BRASSOLINAE 
--HAETERINAE 



Fig. 327. A proposed phylogeny of the Satyridae showing the relation- 
ships of the tribes. The time scales are very approximate and conform to the 
hypotheses proposed in the section on evolution and zoogeography. For details, 
see text. 



LEE D. MILLER 145 

The Neotropics. — The Neotropics, as broadly and somewhat incor- 
rectly defined to include Antarctic, temperate and tropical regions 
of South America, show a satyrid fauna composed of six tribes which 
are members of four subfamilies. Five of these tribes are endemic, 
and four of them — Haeterini, Brassolini, Antirrhini and Biini — are 
the most primitive in the family. The fifth endemic tribe, the Pro- 
nophilini, is a member of the Satyrinae and most closely related to 
the Holarctic Satyrini. The pronophilines probably reached, during 
the mid-Tertiary, South America and proliferated there. The Eup- 
tychiini are exclusively American satyrids most closely allied to the 
Ypthimini. They were probably derived from a common stock with 
the Ypthimini, introduced into the Neotropics and proliferated there 
contemporaneous with the Pronophilini. The euptychiines, while 
entirely American and overwhelmingly Neotropical, cannot be con- 
sidered an endemic tribe because seven species are found in the 
Nearctic, as far north as southern Canada. 

In summary the Neotropical satyrid fauna appears to have been 
derived from two sources. The Haeterinae, Brassolinae, Antirrhini 
and Biini evolved in tropical America from the basic satyrid stock 
which either arose in tropical America from a basic nymphaloid stock 
that was already there or evolved from such a stock that invaded 
tropical America during the Cretaceous. The Neotropics were then 
reinvaded, probably in the Oligocene, perhaps even in the Eocene, 
by two basically Paleotropical groups, the Pronophilini (derived 
from a Satyrini-section stock) and the Euptychiini (from a Ypthimini- 
section stock). There has been no extensive immigration in recent 
times from the Nearctic; indeed, most recent movement in the Amer- 
icas of the Satyridae has been northward from the Neotropics. 

The Nearctic. — The Nearctic fauna has been derived primarily 
from the Neotropics and the Palearctic. Seven tribes are represented 
in North America, none endemic, and most species are very recent 
arrivals, many during the Pleistocene. By far the majority of the 
species in the Nearctic are assignable to Holarctic tribes or genera. 
A few of these, such as Neominois ridinsii, Enodia portlandia and 
creola, Satyrodes eurydice, Coenonympha haydenii and Cercyonis 
species, appear to be pre-Pleistocene arrivals; these butterflies prob- 
ably arrived in North America in the Miocene or Pliocene, perhaps 
even earlier. The remainder of the genera arriving from the Palearctic 
— Oeneis, most Coenonympha, Erebia — belong to a basically arctic- 
MEM. AMER. ENT. SOC, 24 



146 THE SATYRIDAE 



Table 7 



The taxonomic composition of the world satyrid fauna. a= single species; 
b = 2 to 5 species; c = 5 to 10 species; d = more than 10 species). 



Taxon 


o 

'a, 
o 
u> 

o 

u 

z 


u 
o 


o 
ca 

a 

Oh 

W 




c 

— 

6 

C 


c 
•— 
< 


o 

< 

o 
H 


OS 

_o 

< 

d 

6 
m 
H 


a 
o 

Hi 

a 

60 

a 


Haeterinae 


d 


















Brassolinae 


d 


















Biinae 


d 




a 




d 


a 


c 


b 


b 


Antirrhini 


d 


















Biini 


a 


















Melanitini 






a 




d 


a 


c 


b 


b 


Manataria 


b 


















Elymniinae 




b 


d 


C 


d 


d 


d 


d 


d 


Lethini 




b 


d 


c 


d 




a 


b 




Zetherini 






a 




b 










Elymniini 










d 


b 


b 


b 


a 


Mycalesini 






c 




d 


d 


d 


c 


d 


Eritinae 










b 










Ragadiinae 






b 




c 










Satyrinae 


d 


d 


d 


d 


d 


d 


d 


d 


a 


Hypocystini 












d 








Ypthimini 






d 


b 


d 


b 


d 


d 


a 


Palaeonympha 






a 














Euptychiini 


d 


c 
















Coenonymphini 




b 


d 


d 












Maniolini 




b 


d 


d 












Erebiini 




c 


d 


d 












Dirini 
















c 




Pronophilini 


d 


a 
















Satyrini 




d 


d 


d 


b 










Melanargiini 






d 


d 












Endemic tribes 


5 







1 


2 


1 





1 






alpine fauna and probably date from the Pleistocene: Coenonympha 
mixturata found in the bogs of Alaska and the Yukon is virtually 
inseparable from the Siberian population and must be a post-Pleisto- 
cene immigrant. The tribes derived from the Palearctic are the 
Lethini, Coenonymphini, Maniolini, Erebiini and Satyrini. Two 



LEE D. MILLER 147 

tribes, the Pronophilini and Euptychiini, have entered the Nearctic 
from the Neotropics. The first of these tribes is represented by a 
single species, Gyrocheilus patrobas, which reaches southern Arizona. 
This invasion is probably of rather recent origin, perhaps post- 
Pleistocene, though it is difficult to assign dates to an invasion of a 
single species. The Euptychiini have left several species in North 
America, and one group — the Megisto cymela complex — has become 
rather distinct from its Neotropical progenitors. The cymela-gvoup 
probably arrived well before the Pleistocene, whereas the other species 
in the Euptychiini are closely allied to tropical members and may 
have arrived during an interglacial. 

The Palearctic. — The importance of the Palearctic, while it was 
not the place of the origin of the Satyridae, nevertheless cannot be 
overstressed with respect to subsequent evolution. Eleven tribes, 
representing four subfamilies, are found in this region, although the 
"true" Palearctic fauna is composed only of the Elymniinae and the 
Satyrinae. The other two subfamilies, the Biinae and the Ragadiinae, 
enter the Palearctic only in its southeastern part, were derived from 
the Indo-Malayan region and need be mentioned only in passing. 
The Elymniinae are represented by the tribes Lethini, Zetherini 
and Mycalesini — the last two being found just in the southeastern 
part of the area and also properly are part of the Indo-Malayan fauna. 
The tribe Lethini is represented chiefly by the Pararge-senes — more 
boreal in its distribution than other lethine groups — which probably 
evolved in the mid-Tertiary and became cold-adapted. The inter- 
esting link between the Lethini and Mycalesini, Mandarinia regalis, 
is found in temperate China, suggesting that the latter tribe may once 
have been found much farther north than it is today. 

The Palearctic is the metropolis of most of the tribes of the 
Satyrinae, with six represented. Five of these tribes — Satyrini, Me- 
lanargiini, Coenonymphini, Maniolini and Erebiini — probably arose 
in the Palearctic itself in response to progressive drying, with the 
increase of grasslands, during the early Tertiary. Only the Melanar- 
giini are truly endemic, all other tribes having expanded into the 
Nearctic during the later Tertiary or Quaternary. The sixth satyrine 
tribe, the Ypthimini, is primarily Paleotropical, and the Palearctic 
ypthimines must have been derived from the tribes. One group within 
the ypthimines, the Callerebia-series, is exclusively Palearctic and 
probably arose from the ypthimine stem about the same time as the 

MEM. AMER. ENT. SOC, 24 



148 THE SATYRIDAE 

other tribes mentioned above. All members of the Callerebia-sevies 
now are found under the stringent ecological conditions of the central 
Asian mountains; none has invaded the main part of the region. 

In sharp contrast with the Nearctic, the Palearctic has been very 
important in the later evolution of the Satyridae. Just why the Pale- 
arctic should have played such a role in the evolution of the Satyrinae 
is a puzzle: there were certainly vast savannas in the Nearctic during 
the Tertiary (Darlington, 1957), which proved to be the route of 
dispersal of the forerunners of the Pronophilini and Euptychiini and 
thus must have been occupied during the Tertiary. The primitive 
forms of the five Holarctic satyrine tribes are all Palearctic, not 
Nearctic. The evidence for these tribes having arisen in the Old 
World is overwhelming. The only tribe which is a candidate for 
arising in part in the Nearctic is the Satyrini: Neominois ridingsii is 
a rather primitive species, most nearly related to the Asian Karanasa. 
The great proliferation of the Satyrus-series (Satyrini) in central 
Asia argues strongly for its being considered the "birthplace" of the 
tribe. In all fairness, however, it must be pointed out that North 
America was the place of the greatest Tertiary development of the 
horses and camels (Darlington, 1957: 352-354), yet no native 
species of either group is presently found in North America. Such 
an evolutionary pattern may explain the satyrids — evolution of some 
stocks in the New World, spread of these stocks into the Old World, 
extinction of the Nearctic fauna and reinvasion and repopulation of 
the Nearctic by Palearctic groups. In the absence of a fossil record, I 
must accept the neontological evidence that the satyrine tribes evolved 
in the areas of their present greatest density and diversity. Parallels 
exist: Simpson (1953: 51-52) mentions the New World oreodonts 
and pronghorn "antelopes" and the Old World giraffes. None of 
these groups left the hemisphere of their origin. 

The Indo-Malayan region. — This area, bounded roughly by India, 
the Philippines and Wallace's line, has been likened to the "Garden 
of Eden" by generations of zoogeographers. Its satyrid fauna is 
strong only in moderately primitive groups. In all, nine tribes of 
five subfamilies are represented in the Indo-Malayan fauna. This is 
the area of the best development of the Ragadiinae and Eritinae, and 
these are to be considered truly endemic to the region. The melani- 
tines (Biinae) are best represented in the Indo-Malayan fauna and 
probably arose there or arrived at an early date. 



LEE D. MILLER 149 

The Elymniinae have reached their highest development in this 
region, where the four tribes are best represented; all probably arose 
there. Those genera tending to connect tribes, with the exception of 
Mandarinia, are found on the Malay Peninsula. The bifurcation of 
the Mycalesini into naked and hairy eyed species took place in this 
region, and both groups are still found throughout the Indo-Malayan 
region. These have spread and provide some rather interesting data 
in the analysis of the African-Malgache fauna. Certain considera- 
tions, outlined in the systematic revision, led to the conclusion that 
the Lethini are the most primitive elymniines. Most lethines have 
hairy eyes, and for this reason the hairy-eyed condition is considered 
primitive in the Mycalesini (see also the discussion under the Mal- 
gache fauna, which follows). 

The Satyrinae are represented by two tribes, the Satyrini and 
Ypthimini. The former is a secondary immigrant from the Palearctic, 
only entering the Indo-Malayan region along its northern end. The 
Ypthimini, however, probably arose in this region and apparently 
replaced its immediate relatives — forms closer to the present-day 
Hypocysta-series (Hypocystini), which is now restricted to the Aus- 
tralian region east of Wallace's line. The ypthimines spread during 
the early Tertiary into Africa and the New World, giving rise to the 
Euptychiini of the latter region. 

In summary, due to the prevalence of the Melanitini, Elymniinae 
and intermediate genera in this region, the Indo-Malayan region is 
considered to have given rise to the elymniines. The Eritinae and 
Ragadiinae also arose in this area. The Indo-Malayan region did 
not play an important role in the later evolution of the Satyrinae: the 
focus of that subfamily shifted to the Palearctic. 

The Australian Region (including the Pacific Islands). — Faunistically 
the Australian region, including Australia, New Guinea, New Zea- 
land and the nearby islands, is one of the most interesting. It is 
distantly related to the Indo-Malayan region, but substantial inter- 
change of faunal elements between these regions occurred only in the 
distant past. Simpson (1961b: 443) has dated the extensive immi- 
gration of the marsupials into Australia as Cretaceous or Paleocene, 
and it was probably at that time that much of the regional butterfly 
fauna arrived. Elsewhere Simpson (1961b: 435-436) shows that 
several waves of rodent groups arrived in the Australian region from 

MEM. AMER. ENT. SOC, 24 



150 THE SATYRIDAE 

the Indo-Malayan from Miocene time to the present. All the tribes 
represented in the Australian region have also penetrated the Pacific 
islands, over water, hence the islands are considered along with the 
continent. 

Just one tribe is endemic in this region, the Hypocystini. The 
metropolis of the Xenica-series is Australia, with some aberrant spe- 
cies in New Zealand. All the satyrids in New Zealand were derived 
long ago from the Australian ones. The Hypocysta-series is best 
developed in New Guinea and has spread into the Pacific islands. 
The other satyrine tribe represented in this region is the Ypthimini, 
the members of which are highly modified, particularly the Pacific 
island genus Xois. The Ypthimini, as shown by their expansion into 
Africa and the Americas, was a fairly plastic group. 

A single Melanitis is found throughout the region in question, 
M. leda, a widespread species distributed west to the westernmost 
part of Africa. The same may be said about those Elymniini which 
occur on these islands; all are widespread species. The arrival of 
Melanitis and Elymnias must have been a recent event since they are 
little differentiated from their continental progenitors. The Myca- 
lesini of the region reflect the probably early spread of this tribe, since 
it also reached Madagascar. 

In summary, the satyrid fauna of the Australian region was 
derived from the Indo-Malayan region, and this derivation very 
early, resulting in a highly distinctive fauna. If the time schedule 
for the marsupials is valid for the satyrids, most of the groups reached 
the region in late Cretaceous to earliest Tertiary times. This indi- 
cates that the Hypocystini of the most evolved subfamily, Satyrinae, 
were in existence by the earliest Tertiary. A few strong flying species, 
such as M. leda, probably arrived over water very recently. 

The Ethiopian Region. — This region includes both the tropical part 
of Africa and temperate South Africa. These are faunistically very 
different areas, but they are so intimately related that they must be 
considered together. 

There is a single endemic African satyrid tribe, the Dirini. This 
tribe, which has morphological affinities with both the Elymniinae 
and the Satyrini-section of the Satyrinae, is restricted to South Africa. 
Several interpretations are possible for the Dirini: (1) the Satyrinae 
arose in Africa, (2) the Dirini were exterminated in all places but 



LEE D. MILLER 151 

South Africa or (3) the Dirini stock was isolated in South Africa 
with the expansion of the tropical rainforest belt into eastern Africa, 
and it evolved there. The first alternative is highly improbable, as 
I have shown in the discussion of the Hypocystini in the systematic 
revision, but either (2) or (3) might be possible. In the absence of 
a fossil record we shall never really know. The only other tribe of 
the Satyrinae represented in the African fauna is the Ypthimini. The 
ypthimines of tropical Africa are closely related to the Indo-Malayan 
ones, whereas most of the South African ypthimines belong to the 
more or less endemic Melampias-series, one member of which is also 
found on Madagascar. It seems likely that the Melampias-series 
were early invaders of Africa — contemporaneous with the invasion 
of the Dirini — which differentiated and were then forced south and 
were replaced in tropical Africa by the ypthimines that arrived later. 

The Lethini show a similar pattern. In southern Africa two 
very primitive lethines, the A eropet es-series, are found which prob- 
ably are remnants of an early fauna. The East and South African 
genus Aphysoneura is much more like the Indo-Malayan lethines, 
and it was probably a later faunal addition. The Mycalesini have 
made a significant invasion of Africa. All the species on the conti- 
nent have naked eyes, unlike those on Madagascar, and probably 
represent a later immigration, contemporaneous with the arrival of 
the "conventional" ypthimines and Aphysoneura. The mycalesines 
will be considered in further detail in the discussion of the fauna of 
Madagascar. The elymniines and the melanitines of Africa are rep- 
resentatives of the most widespread groups and almost certainly 
arrived from the Indo-Malayan region, probably fairly recently: 
frequently the African members are conspecific with their Indo- 
Malayan counterparts. 

In summary, Africa appears to have been populated more than 
once from the Indo-Malayan region. The tropical African satyrid 
fauna looks more like a depauperate Indo-Malayan fauna than any- 
thing else, but that of temperate Africa has evolved significantly and 
is quite distinct. Probably the South African endemic groups rep- 
resent the "old" African fauna of the early Tertiary, whereas that 
of tropical Africa is younger, perhaps dating from the Oligocene or 
even the Miocene (Darlington, 1957: 590). Such species as Mela- 
nitis leda probably arrived in tropical Africa much more recently. 

MEM. AMER. ENT. SOC, 24 



152 THE SATYRIDAE 

Madagascar. — There can be little doubt that the colonization of 
Madagascar was from Africa, but mostly so long ago that the species 
bear little resemblance to species now found on continental Africa. 
The Melanitini and the Elymniini of this island, on the other hand, 
are conspecific with their mainland counterparts, as is the only mem- 
ber of the Satyrinae, a species of Ypthimini: all these butterflies are 
probably recent introductions, no doubt by waif dispersal. 

The most interesting satyrids in Madagascar, from a zoogeo- 
graphic standpoint, are the Mycalesini — the dominant tribe on the 
island. As noted before, both naked- and hairy-eyed mycalesines 
occur in the Indo-Malayan regions, whereas those from continental 
Africa have naked eyes. The species found in Madagascar, without 
exception, have hairy eyes. One would be tempted at first glance to 
postulate that the Malgache mycalesines were derived directly from 
the Indo-Malayan ones. This is precisely why the mythical continent 
of "Lemuria" was postulated: a land bridge to account for the similar 
Malgache and Indo-Malayan lemurs; the Madagascar genera are less 
like continental African genera than any of the continental genera 
are like each other. In the mycalesines, dissimilarities indicate a long 
isolation of the Malgache fauna. For this reason it seems more 
likely that the mycalesines migrated into Africa very early (Creta- 
ceous or Paleocene?), and that stock which made the first immigra- 
tion was hairy-eyed, as is characteristic of primitive elymniines. Fur- 
thermore, the Malgache mycalesines have a less clubbed female 
foretarsus than do the continental species. These mycalesines were 
contemporaneous with the Dirini, Melampias-series of the Ypthimini 
and the Aeropetes-series of the Lethini and formed part of the "old" 
African satyrid fauna. Madagascar received its mycalesine fauna 
from this very early invasion. Just when this occurred is problematic: 
Matthew (1915: 204-205) thought that Madagascar and Africa 
were never physically connected, and Darlington (1957: 535) be- 
lieved that the Malgache fauna may have begun to accumulate in the 
Oligocene. In any event, the Malgache mycalesine fauna is probably 
of early Tertiary age. Climatic changes or pressure of competition by 
the later mycalesine immigrants — which did not reach Madagascar — 
replaced the hairy-eyed mycalesines on mainland Africa, leaving the 
morphologically primitive Malgache fauna completely isolated geo- 



LEE D. MILLER 153 

graphically from their nearest relatives in the Indo-Malayan region. 
Such invasion and subsequent extermination of geographically inter- 
mediate populations is well known among fossil vertebrates (Dar- 
lington, 1957: 102-121; 159-172; 206-230). 

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Rhopalocera. Satyridae). Berliner Ent. Zeits. 50: 43-141. 

. 1906. Monographische Bearbeitung der Gattungen Lasiophila 

Felder, Daedalma Hew., Catargynnis Rober, Oxeoschistus Butl., Prono- 
phila Westw., Corades Doubl. Hew. (Lepidoptera Rhopalocera. Saty- 
ridae). Mit Begriindung neuer Gattungen und einer Anzahl Neubesch- 
reibungen. Berliner Ent. Zeits. 51: 101-233; ill. 

Tutt, I. W. 1896. British butterflies, being a popular handbook for young 
students and collectors. London: 476 pp.; ill. 

Verity, R. 1920. Contributo alle ricerche sulla variazione e la distribuzione 



LEE D. MILLER 163 

dei Lepidotteri in Italia: Zygaenides, Grypocera e Rhopalocera del Mas- 
siccio delle Mainardi. Boll. Lab. Ziol. Portici 14: 33-61. 

1953. Le farfalle diurne d'ltalia. Volume quinto. Divisione 



Papilionoidea. Sezione Nymphalina: famiglia Satyridae. Florence, 

Italy, Casa editrice Marzocco: xix + 354 pp.; ill. 
Wallengren, H. D. J. 1853. Skandinaviens Dagfjarilar — Lepidoptera 

Scandinaviae Rhopalocera, deposita et descripta. Malmo. Sweden: xx 

+ 280 pp. 
. 1857. Kaffernlandets Dag-Fjarilar, insamlade aren 1838-1845 

af J. A. Wahlberg. K. Svenska Vet.-Akad. Handl. 2(4) : 1-55. 
. 1858. Nya Fjaril-slagten. K. Svenska Vet.-Akad. Handl. 15: 



75-84, 135-142. 

Warren, B. C. S. 1936. Monograph of the genus Erebia. London, Trustees 
of the British Mus. (Nat. Hist.): v-vii + 407 pp.: ill. 

. 1947. Some principles of classification in Lepidoptera with spe- 
cial reference to the butterflies. The Entomologist 80: 208-217. 235- 
241. 

Waterhouse, G. A. and G. Lyell. 1914. The butterflies of Australia. . . 
Sydney, Angus and Robertson: iv-vi + 239 pp.; ill. 

Watkins, H. T. G. 1925. New Callerebias (Lepidoptera. Satyridae). Ann. 
Mag. Nat. Hist. (9)16: 233-237. 

. 1928. New satyrid butterflies. Ann. Mag. Nat. Hist. (10)1: 

615-618. 

Westwood, J.O. 1841. //; Humphreys, H. N., and J. O. Westwood, British 
butterflies and their transformations, ed. 1. London: xii + 138 pp.; ill. 

Weymer, G. 1890. in Weymer, G., and P. Maassen, Lepidopteren gesam- 
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lin, Priv. publ.: vi + 182 pp. 

. 1909. Eine neue Gattung und zwei neue Arten aus der Familie 

der Satyriden. Ent. Zeits. Stuttgart 23: 163-165. 

1910-1912. Satyridae. in Seitz. A., Die Grossschmetterlinge 



der Erde, vol. 5 (Die Amerikanische Tagf alter). Stuttgart, Alfred Ker- 

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a new genus and species of rhopalocerous Lepidoptera from South India. 

Jour. Asiatic Soc. Bengal 49(2): 248-250. 
Zeller, P. C. 1850. Zwei neue Tagf alter. Stettiner Ent. Zeitg. 11: 308-313. 



MEM. AMER. ent. soc, 24 



164 



THE SATYRIDAE 



Index to Taxa 
The numbers in bold face type indicate that the taxon is figured 
on that page. Those names preceded by a $ mark are preoccupied 
names. 



abdelkader Pierret, 123 

abeona Donovan, 81 

abretia Capronnier, 92 

achanta Donovan, 79, 81 

acraeida Butler, 118 

Acrophtalmia Felder & Felder, 67, 68, 69 

" Acrophthalmia" Felder & Felder, 69 

Acropolis Hemming, 69 

actaea Linne, 121, 123 

actoriaena Linne, 32-34 

Ad mi ratio Hemming, 58, 62, 63 

adolphei Guerin, 63 

adoptiva Weymer, 31, 32 

aega Fabricius, 132 

aegeria Linne, 48, 50 

aello Hiibner, 124 

Aeropetes Billberg, 43, 44, 46. 47, 48, 52 

Aeropetes-series, 40, 41, 42. 43. 47, 72, 

137, 139, 151, 152 
aethiops Esper, 105 
Agapetes Billberg, 126 
Agapetinae, 124 
agondas Boisduval, 56, 57 
Agrusia Moore, 56 
albinotata Butler, 94 
albonotata Godman, 117 
albopunctata Weymer, 117 
alini Bang-Haas, 103, 104 
alope Fabricius, 101 
A Itopedaliodes Forster, 1 1 4 
Amathusia Fabricius, 132 
Amathusiidae, 131 
ambiorix Wallengren, 117 
Amecera Butler, 50 
Amechania Hewitson, 51, 52, 53 
Amphidecta Butler, 110, 111, 113, 114 
amphirhoe Hiibner, 27 
Anadebis Butler, 50 
anaxias Hewitson, 63 
anaxioides Marshall & deNiceville, 63 
Anchiphlebia Butler, 31 
andromacha Hubner, 47 
"andromeda Fabricius", 22 
anthe Ochsenheimer, 123 



antipodum Doubleday, 79, 80 
Antirrhea Hubner, 9, 30, 31, 32 
Antirrhini, 26, 28, 29-32, 34, 35, 36, 37, 

133, 136, 138, 144, 145, 146 
antonia Staudinger, 114 
Antopedaliodes Forster, 114 
Apaturina Herrich-Schaffer, 19 
Aphantopus Wallengren, 95, 96, 97, 98 
Aphysoneura Karsch, 43, 47, 48, 151 
aramis Hewitson, 62 
Aranda Fruhstorfer, 49 
arcesilaus Cramer, 31 
archaea Hubner, 31, 32 
Archeuptychia Forster, 92 
Archondesa Moore, 47 
areolatus Smith & Abbot, 94 
aretliusa Esper, 123 
Arethusana deLesse, 123 
XArge Hubner, 126 
Arge Schrank, 126 
arge Sulzer, 126 
Argeformia Verity, 124, 126 
argent ens Blanchard, 110, 112, 117 
Argestina Riley, 87 
Argus Bohadsch, 49 
% Argus Scopoli, 49 
Argynnina Butler, 75, 77, 80 
Argyreuptychia Forster, 93 
Argyronympha Mathew, 77, 80 
Argyrophenga Doubleday, 74, 75, 78, 79, 

80 
Argyrophorus Blanchard, 109, 110, 112. 

113, 117 
armandi Oberthur, 123 
armilla Butler, 94 
Arpidea Duncan, 31 
artemis Felder & Felder, 68, 69 
ashna Hewitson, 94 
asiatica Oberthur & Houlbert, 126 
asochis Hewitson, 63 
Atercoloratus Bang-Haas, 102, 103 
atreus Kollar, 24 
Auca Hayward, 110, 117 
Aulocera Butler, 123 



LEE D. MILLER 



165 



"aurora Felder", 22 
automedon Cramer, 27 
autonoe Cramer, 94 
autonoe Esper, 123 

buladeva Moore, 47 

baldus Fabricius, 85 

banghaasi Weymer, 94 

bathseba Fabricius, 101 

bega Westwood, 112, 117 

Berberia deLesse, 123 

bhadra Moore, 49 

Bia Hiibner, 29, 30, 32-34, 136 

Bicyclus Kirby, 58. 62, 63 

Biina, 3, 4, 28, 32 

Biinae, 3, 4, 8, 9, 11, 12, 13, 16, 17, 18, 

19, 23, 28-36, 39, 72, 132, 133, 135, 

136, 140, 144, 146, 147, 148 
Biini. 10, 28, 29, 30, 32-34, 133, 144, 

145, 146 
Blanaida Kirby, 49 
blanda Moschler, 93 
Blelogona Felder & Felder, 40, 57, 58, 

59, 60, 61 
Boeberia Prout, 86, 87 
bolanica Marshall, 85 
Bolboneura Godman & Salvin, 16 
brahminus Blanchard, 123 
Brassolidae, 4, 23 
Brassolides, 2, 23 
Brassolina, 3, 4 
Brassolinae, 2, 3, 4, 8, 9, 11, 13, 16, 17, 

18, 19, 23-28, 29, 72, 133, 136, 140, 

144, 145, 146 
Brassolini, 23-28, 133, 144, 145 
Brassolis Fabricius, 24, 25, 26, 27 
Brassolis-series, 25, 26, 27 
Brintesia Fruhstorfer, 121, 123 
briseis Linne, 123 
Bruasa Moore, 54, 56 
butleri Fereday, 79, 80 
byses Godart, 93 

Caerois Hiibner, 8. 30, 31, 32 
caerulea Butler, 92 
Caeruleuptychia Forster, 92 
Caliginae, 3, 4, 23 
Caligo Hiibner, 23, 24, 25, 26, 27 



Caligo-series, 25, 27 
Calislo Hiibner, 113. 114 
Callarge Leech, 51, 52, 53 
Callerebia Butler, 82, 86, 87 
Callerebia-series, 83, 87, 102, 141. 147, 

148 
Callicore Hiibner, 16 
Callinaga Moore, 19 
callipteris Butler, 47 
Callitaera Butler, 22, 134 
Callyphthima Butler, 86 
Calysisme Moore, 61 
Capronnieria Forster, 92 
cassiae Linne, 24 
cassina Butler, 108 
Cassionympha vanSon, 86 
cassiope Cramer, 27 
cassiope Fabricius, 105 
cassius Godart, 86 
Cassus vanSon, 106, 107, 108 
cassus Linne, 107, 108 
Catargynnis Rober, 112, 114 
Catoblepia Stichel, 25, 27 
cecilia Vallantin, 101 
Celebina Fruhstorfer, 61 
Cepheuptychia Forster, 92 
cephus Butler, 92 
Cercyonis Scudder, 99, 100. 101, 120, 

145 
ceto Hiibner, 104 
Charma Doherty, 47 
Chazara Moore, 123 
Cheimas Thieme. 112, 114 
clwiiui Guerin, 85 
"chiliensis C. et R. Felder". 118 
chiliensis Guerin, 118 
Chillanella Herrera, 118 
Chionobas Boisduval, 120, 124 
Chloreuptychia Forster, 92 
clitoris Cramer, 92 
Chonala Moore, 49 
Choranesa Moore, 47 
chorinaeus Fabricius, 31, 32 
Chortobius [Dunning & Pickard], 96, 97, 

98 
christophi Leech, 49 
circe Fabricius, 121, 123 
cirta Felder & Felder, 1 1 6 
Cissia Doubleday, 91, 92 



MEM. AMER. ENT. SOC, 24 



166 



THE SATYRIDAE 



Cithaerias Hiibner, 20, 21, 22 

clarissa Cramer, 92 

clio Weymer, 94 

Clothilda Blanchard, 133 

cluena Drury, 92 

clyte Hiibner, 108 

clytus Linne, 107, 108 

Coelites Westwood & Hewitson, 64, 65, 

66, 67, 72 
Coenonympha Hiibner, 96, 97, 98, 142, 

145 
Coenonymphini, 9, 10, 11, 70, 73, 74, 75, 

81, 89, 95-98, 99, 142, 144, 146, 147 
Coenyropsis vanSon, 86 
Coeruleotaygetis Forster, 93 
Colaenis Hiibner, 19 
constantia Cramer, 35, 36 
Corades Doubleday, 112, 114 
corderoi Dognin, 114 
Corderopedaliodes Forster, 114 
Cosmosatyrus Felder & Felder, 118, 127 
Crebeta Moore, 44, 48, 150 
creola Skinner, 145 
crisia Geyer, 68, 69 
Culapa Moore, 59, 60, 61 
cyclopina Staudinger, 115, 118 
cyllastros Westwood & Hewitson, 24, 28 
Cyllo Boisduval, 36 
Cyllogenes Butler, 30, 35, 36 
Cyllopsis R. Felder, 90, 91, 93 
cymela Cramer, 87, 90, 91, 94, 147 

Daedalma Hewitson, 109, 112, 116 

Dalapa Moore, 58, 59, 61 

Dallacha Moore, 82, 83, 85 

damaris Doubleday, 50 

Danaidae, 15, 19, 55, 130, 134 

Danaus Latreille, 130 

% Dasyomma Felder & Felder, 61 

Dasyomma Macquart, 61 

Dasyophthalma Westwood, 24, 25, 26, 27 

Dasyophthalma-series, 25, 27 

Davidina Oberthur, 119, 120, 121, 122. 

123 
Debis Doubleday & Hewitson, 47 
decorata Butler, 117 
decorata Felder & Felder, 116 
deidamia Eversmann, 48, 50 
dendrophilus Trimen, 47, 48 
dexamenus Hewitson, 61 



diademoides Moore, 50 

Dichothyris Karsch, 58, 62, 63 

Didonis Hiibner, 56 

digna Marshall, 123 

Dingana vanSon, 106, 108 

dingana Trimen, 108 

dinias Hewitson, 112, 116 

Dionana Moore, 47 

Dioriste Thieme, 111, 116 

Dira Hiibner, 106, 107, 108 

Dirini, 9, 10, 12, 41, 71, 72, 73, 102, 

105-108, 119, 124, 139, 143, 144, 146. 

150-151, 152 
Dodonidia Butler, 74, 75, 78, 79, 80 
doraete Hewitson, 116 
dor us Esper, 98 
drepana Westwood, 62, 63 
Drucina Butler, 110, 111, 116 
dntsia Cramer, 61 
drusillodes Oberthur, 61 
Drusillopsis Fruhstorfer, 61 
dryas Scopoli, 123 
Dubierebia Muschamp, 98 
Dulcedo d'Almeida, 20, 22 
dumetorum Oberthur, 48, 50 
duplex Butler, 87 
duponcheli Guerin, 63 
Dyctis Boisduval, 55, 56, 57 
Dynastor Westwood & Hewitson, 25, 26, 

27 
dynsate Hewitson, 47 

Elina Blanchard, 109, 115, 117 

Elina-series, 108, 110, 117 

Elymnias Hiibner, 54, 56, 57, 132, 150 

Elymniina, 3, 4, 38, 54 

Elymniinae, 3, 4, 8, 9, 12, 13, 16, 17, 

18, 19, 23, 26, 29, 38-63, 64, 67, 72, 

135, 138, 139, 140, 144, 146, 147, 149. 

150 
Elymniini, 10, 11, 12, 13, 17, 18, 23, 26, 

39, 40, 41, 42, 52, 54-57, 135, 137, 

139, 144, 146, 150. 152 
Elymniopsis Fruhstorfer, 54, 55, 56, 57 
emma Staudinger, 116 
Enodia Hiibner. 40, 41, 43, 46, 47 
Enodiinae, 3, 4, 38, 40, 41 
Enodiini, 41 
t Enope Moore, 49 
Enope Walker, 49 



LEE D. MILLER 



167 



enyo Hewitson, 112, 114 

Epigea Hiibner, 104 

Epimede Houlbert, 126 

epimenides Menetries, 50 

Epinephele Hiibner, 101. 109 

"Epinephila" auctt., 101 

"Epinephile" auctt., 101 

epiphron Knoch, 105 

epistygne Hiibner, 103, 105 

Erebia Dalman, 74, 78, 80, 102, 103, 

104, 105, 108, 114, 142, 145 
Erebiinae, 101 
Erebiini, 9, 71, 73, 81, 95, 98, 101-105, 

108, 119, 142, 144, 146, 147 
erebioides Felder & Felder, 115, 118 
Erebiola Fereday, 74, 78, 79, 80 
t Erebonwrpha Elwes, 87 
Erebomorpha Walker, 87 
Eretris Thieme, 116 
ericluho Butler, 93 
Ericluhodes Forster, 93 
Elites Westwood, 64, 65, 66 
Eritinae, 4, 8, 9, 12, 16, 17, 19, 39, 64- 

66, 67, 72, 135, 139, 140, 144. 146, 

148, 149 
Eritini, 64-66, 144 
Erycinidia Rothschild & Jordan, 75. 76, 

80 
Eryphanis Boisduval, 25, 26, 27 
esaca Westwood & Hewitson, 56 
Etcheverrius Herrera, 118 
Eteona Westwood, 109, 111, 116 
Ethope Moore, 44, 50 
Eumenis Hiibner, 123 
euphemia Westwood & Hewitson, 76, 80 
Euploea Fabricius, 55 
Euploeamima Holland, 50 
Euptychia Hiibner, 90, 91, 93 
Euptychiini, 9, 70, 73, 74, 81, 87. 88, 

89-95, 98, 99, 140, 141, 142, 144, 145, 

146, 147, 148, 149 
Euptychoides Forster, 93 
euripides Weymer, 117 
europa Fabricius, 45, 48 
eurydice Linne, 46, 49, 145 
eurylochus Hiibner, 24, 27 
Eurytelidae, 38, 54 
eurytus Fabricius, 94 
evadne Cramer, 61 



fagi Scopoli, 123 

Faunis Hiibner, 132 

Faimula Felder & Felder, 118 

fergana Oberthiir & Houlbert, 126 

francisca Stoll, 60, 61 

fumata Butler, 95 

fuscum Felder & Felder, 60, 61 

galathea Linne. 125, 126 

t Gallienia Oberthiir, 63 

G are ris Moore, 61 

"Geirocheilus" Holland, 116 

Geitoneura Butler, 78, 79, 80 

germainii Felder & Felder, 118 

geticus Esper, 98 

glacial is Moll, 124 

Gnophodes Westwood, 36 

Godartiana Forster, 93 

Gorgo Hiibner, 102, 104 

gotama Moore, 63 

gracilis Rothschild & Jordan, 76, 80 

Gyrocheilus Butler, 109, 110, 111. 112. 

113, 116 
gyrtone Berg, 1 17 

Haetera Fabricius, 20, 21, 22, 134 
Haeterinae, 3, 4, 7, 8, 9, 10, 11, 12. 13. 

15, 16, 17, 18, 19-22, 23, 29, 72, 132. 

133, 134, 136, 140, 144, 145, 146 
Haeterini, 19, 20-22, 133, 144, 145 
Halimede Oberthiir & Houlbert, 126 
haiimede Menetries, 126 
Hallelesis Condamin, 63 
Hamadryopsis Oberthiir, 61 
"Hanifa" auctt., 47 
hanifa Nordmann, 123 
Hanipha Moore. 47 
Hariina Moore. 47 
Harjesia Forster, 93 
Harsiesis Fruhstorfer, 80 
haydenii Edwards, 145 
Haywardella Herrera, 116 
Haywardina Aczel, 93 
t Haywardina Forster, 93 
hedemanni R. Felder, 91, 93 
helmsii Butler, 79, 80 
Hemadera Moore, 85 
Henotesia Butler, 63 
hercyna Hiibner, 36, 37 



MEM. AMER. ENT. SOC, 24 



168 



THE SATYRIDAE 



hermes Fabricius, 93 

Hermeuptychia Forster, 93 

Hermianax Fruhstorfer. 50 

Hermias Fruhstorfer, 42, 44, 47 

hermione Linne, 123 

herse Cramer, 93 

hesione Cramer, 62 

hesione Sulzer, 94 

Hetaerina, 3, 4, 19, 20 

Heteronympha Wallengren, 75, 77, 81 

Heteropsis Westwood, 58, 62, 63 

hewitsonii Doumet, 62, 63 

hilda Westwood, 50, 51 

himachala Moore, 50 

Hipio Hiibner, ,30, 35, 36 

Hipparchia Fabricius, 119, 123 

Hipparchiadae, 15, 69, 119 

Hipparchioid.es Butler, 81 

hippia Cramer, 87 

hobartia Westwood & Hewitson, 80 

Homoeonympha Felder & Felder, 109, 

118 
Houlbertia Oberthiir, 63 
huebneri Felder, 123 
huebneri Kirby, 85 
humilis Felder & Felder, 118 
hyagriva Moore, 85 
Hyalodia Jordan, 75, 80 
hygeia Hewitson, 80 
hypaesia Hewitson, 22 
"hyperanthus" auctt., 98 
hyperantus Linne, 97, 98 
hyperbius Linne, 86, 87 
hypermnestra Linne, 56, 57 
Hypocysta Westwood & Hewitson, 75, 

76, 80 
Hypocysta-series, 12, 74, 76-77, 78, 80, 

81, 149, 150 
Hypocystini, 7, 9, 11, 12, 57, 70. 72, 73. 

74-81, 102, 105, 108, 110, 119, 124, 

139, 140. 141, 144, 146, 149, 150, 

151 
Hyponephele Muschamp, 101 

I data deLesse, 101 

Idiomorphus Chaudoir, 63 

i Idiomorphus Doumet, 63 

t Idioneura Felder & Felder, 118 

Idioneura Selys, 1 1 8 



Idioneurula Strand, 110, 113, 115, 118 

incerta Hewitson, 51, 53 

Indalasa Moore. 61 

inga Fruhstorfer, 61 

inica Hewitson, 85 

Ithomiidae, 9, 12, 14-15, 16, 19, 130, 

131-132, 134 
itonia Hewitson, 85 

jalaurida deNiceville, 49 
janardana Moore, 61 
janira Linne, 101 
J at ana Moore, 61 
jurtina Linne, 100, 101 
jynx Hiibner, 57 

Kabanda Moore, 61 

kalinda Moore, 87 

Kallima Doubleday, 19 

Kanetisa Moore, 123 

Karanasa Moore. 119, 123, 148 

Kerrata Moore, 47 

Kirinia Moore, 4.3, 50 

Kirrodesa Moore, 47 

klugii Guerin, 79, 80 

Kolasa Moore, 82, 85 

Lachesis Oberthiir & Houlbert, 126 

lachesis Hiibner, 126 

lais Cramer, 57 

lais Fabricius, 57 

Lampides Hiibner, 130 

Lamprolenis Godman & Salvin, 72, 74, 

75, 76, 77, 80, 139 
lanaris Butler, 47 
languida Butler, 94 
Lasiommata Westwood, 50 
Lasiophila Felder & Felder, 110, 116 
lathionella Westwood, 81 
latipicta Fruhstorfer, 50 
leaena Hewitson, 118 
leda Gerstacker, 87 
leda Leech, 126 

leda Linne, .35. 36, 136, 150, 152 
Ledargia Houlbert, 124, 126 
lemur Schrank, 101 
leonata Butler, 111, 116 
leprea Hewitson, 81 
Leptoneura Wallengren, 108 



LEE D. MILLER 



169 



leptoneuroides Felder & Felder, 118 

Lepiotes Scudder, 130 

Lethe Hubner, 16, 41, 43, 44. 45, 47, 48 

Le//je-series, 42, 43, 47-49, 137 

Lethinae, 3, 4, 38, 40, 41 

Lethini, 28, 34, 37, 38, 39, 40-51, 52. 54. 
57, 64, 72, 74, 105, 135, 137, 138- 
140, 144, 146. 147, 149, 151, 152 

leucoglene Felder & Felder, 1 1 8 

libye Linne, 94 

Libythea Fabricius, 130 

Libytheidae, 130 

ligea Linne, 103. 104 

Limenitinae, 52 

Use Hemming. 56. 57 

Loesa Moore, 61 

Lohana Moore, 85 

Lohora Moore, 61 

Lopinga Moore, 44, 48, 50 

lorquinii Felder & Felder, 46, 49 

lowi Doubleday & Hewitson, 50 

Loxerebia Watkins, 82, 87 

Lycaena Fabricius, 74 

Lycaenidae, 130 

lycaon Rottemburg, 101 

Lycoreinae, 133 

Lyela Swinhoe. 95, 96, 98 

Lymanopoda Westwood, 109, 118 

Lymanopoda-scnes, 108, 110, 113, 114, 
118, 127 

lymessa Hewitson, 81 

macmahoni Swinhoe, 98 

madura Horsefield, 66 

Magneuptychia Forster, 94 

t Magula Fruhstorfer, 44, 49 

Magula Scudder, 49 

maianeas Hewitson, 63 

malsara Moore, 49 

mahahda Butler. 61 

Manataria Kirby, 10, 18. 34. 36. 37, 38. 

41, 146 
Mandarinia Leech, 41, 43, 46, 50, 51, 

72, 74, 139. 149 
Mandarinia-stries. 42, 43, 50 
Manerebia Staudinger, 110, 111, 115. 

118 
Maniola Schrank 2, 99. 100. 101, 109 
Maniolidi, 98 



Maniolinae, 69, 98 

Maniolini. 9, 71, 73, 74, 81, 89, 98- 

101, 109, 120, 142, 144, 146, 147 
margaretae Elwes, 47 
Market Hubner, 104 
marshallii Wood-Mason, 36 
Martanda Moore, 58, 59, 61 
Mashuna vanSon, 83, 84, 85 
mashuna Trimen, 84, 85 
masoni Elwes, 49 
Masoura Hemming, 58, 63 
masoura Hewitson, 59, 63 
inedus Fabricius, 60, 62 
medus Schiffermiller, 104 
Medusia Verity, 102, 104 
megera Linne, 50 
Megeuptychia Forster, 94 
Megisto Hubner, 91, 94 
Melampias Hubner, 82, 86, 87 
Melampias-series, 83, 84, 86-87, 141. 

151, 152 
Melanargia Meigen, 4, 11, 16, 124, 125, 

126 
Melanargiinae, 124 
Melanargiini, 5, 9, 10, 71, 73, 105. 119. 

124-126, 143, 144, 146, 147 
Melania Lamarck, 123 
t Melania Sodoffsky, 123 
Melanitini, 10, 18, 28, 29, 30, 34-36, 37, 

39, 41, 135, 136, 138, 144, 146, 149, 

152 
Melanitis Fabricius, 16, 30, 35, 36, 150 
Melynias Moore, 54, 55, 57 
menander Drury, 22 
Meneris Westwood, 47 
mermeria Cramer, 91, 95 
merope Fabricius, 81 
Mestra Hubner, 11, 16 
mestra Hewitson, 62 
Mimadelias Moore, 54, 57 
minerva Fabricius, 49 
Minois Hubner, 99, 119, 120, 123 
Mint ha vanSon, 108 
mintha Geyer, 107, 108 
mirabilis Staudinger, 118 
mixturata Alpheraky, 95, 146 
mnasicles Hewitson, 60, 61 
mollina Hubner, 91, 92, 93 
monachus Blanchard, 117 



MEM. AMER. ENT. SOC, 24 



70 



THE SATYRIDAE 



Moneuptychia Forster, 94 

Monotrichtus Hampson, 61 

monticolens Butler, 118 

moorei Felder, 61 

Morphidae, 2, 3, 14-15, 16, 19, 23, 131, 

132, 134 
Morpho Fabricius, 15, 132 
Muscopedaliodes Forster, 1 1 6 
muscosa Thieme, 116 
Mycalesini, 11, 12, 13, 33, 39, 40, 52, 

54, 57-63, 72, 74, 75, 128, 135, 138, 

139, 144, 146, 147, 149, 150, 151, 152- 

153 
Mycalesis Hubner. 42, 46, 58, 60, 61, 

63, 139 
Mycalesis group, 67 
mycalesis Felder & Felder, 61 
mycalesoides Felder, 94 
Mydosama Moore, 60, 61 
Mygona Westwood, 116 
mynois Hewitson, 61 
myops Staudinger, 98 
Myrtilus deNiceville, 61 
mystes deNiceville, 61 

Nadiria Moore, 85 

napoleon Westwood & Hewitson, 27 

narasingha Moore, 85 

nareda Kollar, 85 

Narope Westwood & Hewitson, 23, 24, 

25, 26, 28 
N a rope-scries, 25, 28 
Nasapa Moore, 62 
natalii Boisduval, 86 
Nebdara Moore, 62 
necys Godart, 93 
Neita vanSon, 87 
neita Wallengren, 87 
Nelia Hayward, 109, 117 
Nemetis Moore, 49 
nemyroides Blanchard, 117 
Neocoenyra Butler, 87 
Neohipparchia deLesse, 123 
Neomaenas Wallengren, 109, 117 
Neomaniola Hayward, 117 
Neominois Scudder, 119, 120, 121, 123 
Neonympha Hubner, 69, 87, 91, 94 
Neope Butler, 49 
Neorina Westwood, 43, 50, 51 
Neorina-series, 37, 39, 40, 42, 43, 50, 52, 



138 
Neosatyrus Wallengren, 117 
nereis Drury, 21, 22 
nesaea Linne, 57 

Nesoxenica Waterhouse & Lyell, 77, 81 
Ninguta Moore, 49 
Nissanga Moore, 62 
nitida Godman & Salvin, 76, 80 
noma Thunberg, 122, 124 
nothis Westwood & Hewitson, 66 
Nymphalidae, 52, 130 
Nymphaloidea, 7, 8, 19 
Nytha Billberg, 123 

obsoleta Westwood, 118 

Ocaiis Westwood, 94 

occitanica Esper, 126 

Odonata, 131 

oedipe Hubner, 98 

oedippus Fabricius, 97, 98 

Oeneis Hubner, 16, 119, 120, 122, 124, 

142, 145 
Oeneis-series, 73, 120, 121, 122, 124, 

143 
opalina Butler, 87, 88 
opalinus Staudinger, 114 
Opoptera Aurivillius, 25, 27 
Opsiphanes Westwood & Hewitson, 24, 

25, 26, 27 
Oreas Hubner, 22 
Oreina Westwood, 105 
Oreixenica Waterhouse & Lyell, 81 
oreseis Hewitson, 63 
Oressinoma Westwood, 90, 92, 93, 94 
Orinoma Gray, 43, 50 
ornata Rothschild & Jordan, 80 
oroatis Hewitson, 61 
Orsotriaena Wallengren, 40, 57, 58, 59, 

60, 62, 139 
Orthoptera, 131 
Oxeoschistus Butler, 116 

Pachama Moore, 62 
Palaeonympha Butler, 74, 87-89, 146 
pales Philippi, 117 
Pal maris Herrera, 118 
Pampasatyrus Hayward, 117 
Pamperis Heimlich, 127-128 
pamphanis Westwood & Hewitson, 27 
pamphilus Linne, 97, 98 



LEE D. MILLER 



171 



Panarche Thieme, 116 

panda Boisduval, 84, 87 

Pandima Moore, 83, 85 

pandoea Hopffer, 63 

Panyapedaliodes Forster, 1 1 6 

panyasis Hewitson, 1 1 6 

Papilio Linne, 2 

Papilionoidea, 3 

parado.xa Mabille, 62, 63 

"Parage" auclt., 50 

Paralasa Moore, 82, 83, 87 

Paralethe vanSon, 9, 43, 44, 47, 48 

Paramecera Butler, 90, 93, 94 

Parantirrhoea Wood-Mason, 30, 34, 35. 

36. 138 
Parapedaliodes Forster, 116 
"Pararga" auctt., 50 
Pararge Hubner, 42, 43, 45, 48, 50 
Para/ge-series, 39, 42, 43, 44, 46, 49-50, 

137, 139, 147 
Parataygetis Forster, 94 
Paratisiphone Watkins, 81 
Parce Oberthiir & Houlbert. 126 
parce Staudinger, 126 
pare pa Hewitson, 116 
Pareuptychia Forster, 94 
parmenio Boeber, 86, 87 
parmeno Westwood, 36 
Paroeneis Moore, 123 
Paryphthimoides Forster, 94 
Pasiphana deLesse, 101 
passandava Ward, 63 
Patala Moore, 49 
patnia Moore, 62 
patrobas Hewitson, 113, 116, 147 
Pedal iodes Butler, 111, 113, 116 
pegala Fabricius, 100, 101 
pelopea Klug, 123 

penanga Westwood & Hewitson, 56 
penelea Cramer, 94 
penelope Fabricius, 91, 93 
Penetes Westwood & Hewitson, 25, 27 
Penrosada Brown, 113, 118 
Percnodaimon Butler, 74. 78, 79, 80, 81 
periboea Fabricius, 128 
periboea Godman & Salvin, 93 
Periplysia Gerstacker, 83, 87 
phaedra Linne, 123 
phanias Hewitson, 116 



phantoma Fassl. 22 

phares Godart, 94 

X Pharia Fruhstorfer, 69 

Pharia Gray, 69 

Pharneuptychia Forster, 94 

t phegea Fabricius, 57 

Pherepedaliodes Forster, 116 

pheretiades Smith & Kirby, 116 

phidia Linne, 123 

Philareta Moore, 123 

philomela Linne, 84, 85 

pholoe Staudinger, 112, 114 

Phorcis Hubner. 102, 103, 104, 105 

phoronea Doubleday, 117 

X Phryne Herrich-Schaffer, 98 

Phryne Meigen, 98 

phryne Pallas, 97, 98 

Physcaeneura Wallengren, 83, 84, 87 

physcoa Hewitson, 116 

Physcon deNiceville, 63 

Physcopedaliodes Forster, 116 

piera Linne, 21, 22 

Pierelta Herrich-Schaffer. 4, 20, 21. 22. 

134 
Pierellinae, 3, 4, 19, 20 
Pieridopsis Rothschild & Jordan, 75, 76, 

78, 80 
pigmentaria Karsch, 47, 48 
pignerator Butler, 111, 114 
pimplea Erichson, 51, 52, 54 
Pindis R. Felder, 90, 94 
pireta Cramer, 21, 22 
Placilla Moore, 49 
Platypthima Rothschild & Jordan, 75, 

77, 80 
pluto Fereday, 79, 81 
poaoeneis Heimlich. 127 
poesia Hewitson, 111, 116 
polita Hewitson, 22 
poltys Prittwitz, 94 
Polymastus Thieme, 116 
polyxo Godman & Salvin, 117 
portlandia Fabricius, 46, 47, 145 
Posteuptychia Forster, 94 
Posttaygetis Forster, 94 
Praefaunula Forster, 94 
Praepedaliodes Forster, 116 
Praepronophila Forster, 116 
pratorum Oberthiir, 87 



MEM. AMER. ENT. SOC, 24 



172 



THE SATYRIDAE 



Precis Hiibner, 130 

pringlei Sharpe, 47 

Proboscis Thieme, 113, 116 

pronoe Esper, 105 

Pronophila Doubleday & Hewitson, 109, 

111, 112, 116 
Pronophila-seiies, 10, 110, 114-117 
pronophila Felder & Felder, 116-117 
Pronophilinae, 3. 4, 108 
Pronophilini, 7. 9, 10, 71, 73, 102, 105, 

108-118, 119, 127, 140. 143, 144, 145, 

146, 147, 148 

propylea Hewitson, 113, 116 
Pseudeuptychia Forster, 94 
Pseudochazara deLesse, 123 
Pseudodebis Forster, 94 
Pseudohaetera Brown, 20, 22 
% Pseudomaniola Rober, 114 
Pseudomaniola Weymer, 114, 117 
Pseudonympha Wallengren, 82, 87 
Pseiidosteroma Weymer, 116-117 
Pseudotergumia Agenjo, 123 
Psyche Hiibner, 126 
Pry chandra Felder & Felder, 34, 39, 41, 

43, 44, 45, 46, 49, 138 
puerta Hewitson, 116 
pulchra Mathew, 80 
pumilus Felder, 123 
Punapedaliodes Forster, 117 
pusilla Felder & Felder, 118 
Putlia Moore, 47 
Pyronia Hiibner, 100, 101 

Quilaphoetlwsiis Herrera, 117 

Ragadia Westwood, 9, 67, 68, 69 
Ragadiina, 3, 4, 67 

Ragadiinae, 3, 4, 8, 9, 12, 13, 15, 16, 17, 
18, 39, 67-69, 72, 139, 141, 144, 146, 

147, 148, 149 
Ragadiini, 67-69, 144 
Rangbia Moore, 45, 49 
Rareuptychia Forster, 94 
regalis Leech, 50, 51, 57, 147 
Rhaphicera Butler, 45, 46, 50 
Rliaphiceropsis Sharpe, 47 
ridingsii Edwards, 121, 123, 145, 148 
ruscinonensis Oberthiir & Houlbert, 126 
rusina Godart, 24, 27 



Sabatoga Staudinger, 1 1 8 

Sadarga Moore, 63 

safitza Hewitson, 61 

sagitta Leech, 51, 53 

saitis Hewitson, 77, 80 

sallei Westwood & Hewitson, 27 

Samanta Moore, 10, 18, 34, 39, 41, 49, 

138 
sambulos Hewitson, 62, 63 
samio Doubleday & Hewitson, 47 
samius Westwood, 110, 115, 118 
Samundra Moore, 63 
sanatana Moore, 61 
Sarromia Westwood, 109, 118 
Satoa Moore, 58, 63 
satricus Westwood & Hewitson, 45, 50 
saturnus Butler, 93 
Satyrides, 2, 6, 15, 69, 119 
Satyrina, 3, 4 
satyrina Bates, 94 
satyrina Butler, 45, 49 
Satyrinae, 3, 4, 6, 7, 8, 9, 10, 12, 13, 

16, 17, 18, 39, 41, 42, 57, 64, 67, 69- 

126, 136, 139, 140-142, 144, 145, 146, 

147, 148, 149, 150, 151, 152 
Satyrini, 6. 7, 9, 10, 71, 73, 99, 105, 106, 

108, 110, 119-124. 142, 143, 144, 145, 

146, 147, 148, 149 
Satyrini-section, 12, 73. 74, 75, 101, 105, 

108, 110, 140, 141, 143, 145, 150 
Satyrodes Scudder, 43, 44, 46, 49 
Satyrotaygetis Forster, 94 
Satyrus Latreille. 2, 119, 121, 123 
Sarynw-series, 10, 120, 122, 123, 143, 

148 
scanda Kollar, 86, 87 
scanda Moore, 45, 49 
schrenkii Menetries, 49 
Selenophanes Staudinger. 25. 27 
servilaea Wallengren, 117 
sesara Hewitson, 84, 85 
Setodocis Billberg, 128 
Sevanda Moore, 63 
Sicca Verity, 98 
sicelis Hewitson, 47 
sidonis Hewitson, 49 
sihala Moore, 47 
similis Butler, 95 
Sim plica Verity, 105 
Sinarista Weymer. 29, 30, 31. 32 



LEE D. MILLER 



173 



Sinchula Moore, 49 

% Smithia Mabille, 63 

Smithia Milne-Edwards & Haime, 63 

sophorae Linne, 24. 26. 27 

soter Butler, 94 

Spinantenna Hayward, 110, 114. 115, 117 

Splendeuptychia Forster, 94 

squamistriga R. Felder. 94 

statilinus Hufnagel, 123 

stelligera Butler. 1 1 8 

Steremnia Thieme, 1 17 

Steroma Westwood, 109, 112, 113, 114, 

117 
Steromapedaliodes Forster, 117 
Stibonwrpha Butler, 117 
Strabena Mabille, 82, 83. 87 
Stuardosatyrus Herrera & Etcheverry, 

117 
Stygionympha vanSon, 87 
stygne Ochsenheimer. 104 
Stygnolepis Strand, 118 
t Stygnus Felder & Felder, 1 1 8 
Stygnus Perty, 118 
sudra Felder, 61 
sura Westwood, 49 
suradeva Moore, 36 
Suralaya Moore, 63 
syme Hiibner, 27 
Syngea Hiibner, 105 

Taenaris Hiibner, 19, 132 

tagala Felder, 62 

tamatavae Boisduval, 87 

Tanaoptera Billberg, 48 

Tansima Moore, 42, 45. 49 

Tarsocera Butler, 106, 108 

Tatinga Moore, 50 

tauropolis Westwood. 1 1 6 

Taygetina Forster, 94 

Taygelis Hiibner. 90, 91. 92. 95 

Telinga Moore, 63 

Tellervini, 132 

tena Hewitson, 1 14 

tenuisquamosa Joicey & Talbot. 80 

Tetraphlebia Felder & Felder, 118 

thalia Leech, 69 

t thelebe Doubleday, 116 

thelebe Doubleday & Hewitson, 111, 116 

Theope Doubleday & Hewitson, 50 



X Theope Moore, 50 

thibetanus Oberthiir, 50 

Thiemeia Weymer, 1 1 7 

thione Berg, 1 1 7 

Thympia Moore. 82, 85 

tircis Stoll, 98 

Tisiphone Hiibner, 81 

tisiphone Boisduval, 116 

tithone Hiibner, 101 

tithonus Linne, 100. 101 

Tithoreini, 132 

tomasia Butler, 31 

Torynesis Butler, 10, 106, 107. 108 

Triariia Verity, 105 

triarius dePrunner, 105 

tricordata Hewitson, 116 

trimacula Leech, 47 

Triphysa Zeller, 95, 96, 97, 98 

tristigmata Elwes, 47 

tristis Guerin, 115. 117 

Triteleuta Strand. 30, 3 1 

Trophonina Rober, 118 

Truucaefalcia Verity, 105 

tulbaghia Linne. 47, 48 

typhla Westwood, 92, 93, 94 

valentina Cramer, 94 

Vanessa Fabricius, 130 

vanessoides Blanchard, 115. 117 

Vareuplychia Forster, 95 

vasudeva Moore, 57 

verma Kollar, 47 

vigilans Trimen, 87 

Virapa Moore, 63 

virgilia Cramer, 95 

virgo Rothschild & Jordan, 76, 80 

viridicans Weymer, 95 

vitellia Cramer, 56 

waltoni Elwes. 87 
wardii Butler, Callyphthima, 86 
wardii Butler, Henotesia, 63 
weidemeyeri Edwards, 52 
Weymerana Forster. 95 
williamsianus Butler, 117 

Xenica Westwood, 75, 78, 79, 81 
Xenica-scries, 74. 76, 77, 78. 80-81, 108, 
150 



MEM. AMER. ENT. SOC, 24 



174 



THE SATYRIDAE 



Xeniconympha Novicky, 81 

xicaque, Reakirt, 93, 94 

Xois Hewitson, 82, 84, 85, 150 

yama Moore, 49 

"Yphthima" auctt., 85 

yphthima Felder, 95 

Yphthimoides Forster, 95 

Ypthima Hiibner, 82, 84. 85 

Ypthima group, 67 

Ypthimaseties, 82, 83, 84, 141 

Ypthimini, 9, 10, 70, 73, 74, 75, 81-87, 
88, 89, 95, 99, 101, 102, 141, 144, 145, 
146, 147, 149, 150, 151, 152 



Ypthimini-section, 12, 73, 74, 75, 81, 90, 

95, 99, 102, 140, 141, 145 
Ypthimomorpha vanSon, 85 
Ypthimorpha Overlaet, 85 

Zabimia Hewitson, 118 

zangris Hiibner, 114 

Zethera C. Felder, 51, 52, 53, 54 

Zetherini, 10, 39, 40, 41, 52-54, 135, 139, 

144, 146, 147 
zigomala Hewitson, 118 
Zipaetis Hewitson, 75, 77, 78, 80 
Zischkaia Forster, 95 
Zophoessa Westwood, 43, 49 






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